ud in their dishes even when they are fed artificially. Main
(1953) has suggested that so-called specific differences in Australian forms may be
due to differences in the salinity of the medium ; I have made attempts to grow
Triops at salinities near those that Main found in the field, but the animals have
rapidly died each time. The temperature at which the animals were reared might
be expected to have some effect on their form, and so a pure line of T. cancriformis
was grown at a temperature (25-28 C) near their lethal point, but these animals
showed little difference from those grown at normal temperatures.
A REVIEW OF THE NOTOSTRACA 7
Notostraca were grown in the laboratory under very diverse conditions of tempera-
ture, food, and vessel size, but all have remained remarkably uniform in their
morphology. Daphnia under parallel conditions would show very marked morpho-
logical changes, and it is probably safe to assume that environmental factors can
affect the morphology only after a very considerable number of generations — as
salinity affects the form of Artemia salina only after several years (Schmankiewitsch,
1875).
The various characters which have been used in systematics are now examined
separately.
(1) Total size
Samples of adult Notostraca usually contain individuals of very different sizes,
and it is difficult to determine whether there is a normal adult size for any species.
The growth rates of animals in a batch of Triops reared in the laboratory are also
very variable (Fox, 1949) as the pure line of T. cancriformis showed very clearly ;
after a few days growth they varied enormously in size ; 33 individuals were between
2*0 and 3*5 mm. carapace length, while one was only 1*5 mm., and another 5-0 mm.
Spandl (1926) gives growth curves for individual T. cancriformis, maintaining that
those which grow the fastest become the largest, which is what one would expect
if the growth rate depended on the level of nutrition.
The usual adult size for all species seems to be between 15 mm. and 30 mm. in
carapace length, the growth curve flattening out somewhere between these two
figures.
The total size has occasionally been used as a systematic character.
Bowkiewicz (1923) suggests that giant Triops which he saw in Siberia might be a
new species ; large individuals of other species occur up to 40 mm in carapace length
and probably correspond to the giant specimens of Anostraca, which Sellier and
Morice (1946) have shown to be cytologically similar to normal individuals in one
species (Chirocephalus diaphanus).
It is difficult to find a suitable measure of the total size of individuals in order
that relative proportions of endites, etc. may be compared during growth in different
samples. The total length has been shown to be quite useless in preserved specimens
(Barnard, 1929 ; Somme, 1934 ; Linder, 1952) and is difficult to measure in living
animals, which have considerable powers of contraction. The measurements made
on living animals also indicate that there is some increase in length during an instar.
Somme (1934) showed that the median carapace length is more reliable for it is
little affected by the action of the preservative.
It has now been possible to show that the growth of the carapace is isometric and
is therefore a valid measurement of size at all ages (p. n).
Unfortunately, the ratio of carapace length /total length varies between samples
and so this measurement is valid only in comparisons within a sample.
Generally, Lepidurus spp. tend to be smaller than Triops spp. and Lepidurus
arcticus is usually smaller than the rest, but probably more rarely achieves its
maximum size than the non-boreal species.
8 A REVIEW OF THE NOTOSTRACA
(2) Colour
Living Notostraca are frequently brightly coloured, the colours being due to two
main pigments within the body and the brown colour of the exoskeleton.
The internal pigments are haemoglobin, which is present in solution in the blood
(Regnard and Blanchard, 1883), and a dark blue-green pigment which occurs in
connective tissue in many parts of the body ; this pigment is of unknown composi-
tion, but is similar in nature to that which occurs in some Ostracods (Fox, 1955).
The concentration of haemoglobin varies inversely with the oxygen tension of the
medium in which the animal is living (Fox, 1949), and large animals in poorly aerated
conditions may have so dense a concentration that the animal appears to be deep
red in colour.
The blue-green pigment appears to be more abundant in Lepidurus than in Triops
and the animal is sometimes deep green in colour (hence Lepidurus viridis,
Baird). In Triops it may be completely absent, though a mid-dorsal patch on the
carapace and smaller areas on the bases of the thoracic appendages are generally
present. It seems to be more abundant in animals which have grown rapidly and
under good conditions, when it forms a dark marbling on the carapace and is well
distributed elsewhere. The only living specimens of T. australiensis that I have
seen have been without this pigment, but this is unlikely to have any significance.
The newly hatched larvae of both genera may be densely coloured with a carotenoid
pigment, as Fox (1949) reported for Triops cancriformis ; in successive generations
of the same stock the colour of the larvae is very variable and probably depends on
the feeding of the parents ; well fed females usually contain eggs which are pink
with carotenoid, while in starved specimens they are white.
The egg shells of Triops are bright red in colour, for a red haemochromogen is
present in them (Fox, 1955) , this pigment is secreted by the follicle ducts of the
ovary, and is brightest in colour in newly laid eggs, becoming duller after they have
been dried.
A bright violet pigment occurs in the egg shell of Lepidurus arcticus (H. Moore,
personal communication), while other species in this genus have eggs of the same
colour as in Triops.
(3) Body-length, segmentation, and number of appendages.
The carapace of Notostraca is attached only to the head region, and the thorax
and abdomen are completely free from it ; the post-carapace region is divided into
a number of segments or body rings, which have considerable powers of telescoping.
The first eleven segments, of which the first is incomplete dorsally, normally each
bear one pair of appendages ventrally, and together comprise the thorax.
The number of post-thoracic, or abdominal, segments is variable and the series
of appendages is continued along them ; a few segments at the posterior end of the
abdomen bear no appendages.
linder (1952) has analysed a great deal of North American material and has to a
great extent elucidated the relationships between the number of segments, the
number of appendages and the number of apodous segments posteriorly. He con-
A REVIEW OF THE NOTOSTRACA 9
eludes that the number of segments and the number of appendages are the results
of two quite separate growth processes for a number of reasons : the production of
segments and appendages in the larvae proceed at different rates ; the boundaries
of the appendage bearing segments are not complete ventrally ; an aberration in
which the segments are spiral instead of annular does not upset the arrangement of
the appendages ; no correlation can be found between the number of appendages
and the segments bearing them ; and the appendage series may end anywhere
along the length of a segment.
While studying the development of the systematic characters during growth I
found that the larvae of Triops complete their segmentation by the 5th or 6th
instar, but that the series of appendages continue to increase and encroach posteriorly
on to fresh segments until the 8th or 9th instar, after which the number of apodous
segments remains constant. After this the number of appendages may continue to
increase for a few instars but come to occupy no further segments. So after the 9th
instar the number of segments and the number which are apodous may be taken as
fixed in an individual, so that these could validly be used as characters in animals
of more than 3-4 mm. in median carapace length.
The total number of segments varies in Triops from 32-44, and in Lepidurus from
26-34 '> these results are based on the museum material combined with information
from the literature. Throughout this paper the figures given do not include the
telson which is considered to be post-segment al, and incomplete segments are
included in the count.
The figures for Triops show that the variation is continuous throughout its range,
and separation into groups on the number of segments would be quite arbitrary ;
the results for females from all sources illustrates this —
No. of segments . 32 33 34 35 36 37 38 39 40 41 42 43
No. of occurrences . 24 31 19 12 11 17 22 12 13 15 10 7
The bottom row of numbers — the number of occurrences — is obtained by counting
the number of samples in which each segment-number occurs ; this prevents undue
weight being given to large samples, which are not evenly distributed along the range
of variation.
The apparent bimodality of these figures is due to the fact that one species —
Triops cancriformis — has a range of segment variation which covers only a part of
that of the other species. This species occurs in Europe and naturally predominates
in the collections, thus increasing the number of occurrences at the lower end of
the scale. T. cancriformis has a variation of 32-35, while the other species probably
run from 32-43.
By far the greatest number of Lepidurus examined fall within the range of 25-29
segments ; I have seen only two samples with more, L. lynchi Linder and L. batesoni
sp. n. This is in accordance with Linder's findings, for he suggested that there
might be two groups of species within Lepidurus based on the number of segments.
The variation within samples bears out the theory that the number of segments
may be of use in the systematics of Lepidurus but not of Triops ; in the pure line
of T. cancriformis there was a variation of 3 in this character (33-35), and variations
io A REVIEW OF THE iNOTOSTRACA
of 2, 3, or 4 are usual in samples of preserved specimens. In Lepidurus, however,
possibly because there are fewer segments, the variation is smaller, rarely being
more than 2 in each sample, many samples showing none.
The number of appendages confirms the above grouping of Lepidurus ; the shorter
bodied group has from 35-48 appendages, the longer from 39-71 ; these figures are
based on Linder's data with the addition of counts made on the material I have
examined. But in Triops I can find no correlation between the number of legs and
the number of segments, for high numbers of appendages occur in both short and
long bodied specimens, and the converse is also true. In this genus I can find no
significance in the numbers of appendages.
The number of apodous segments is a secondary character depending on the
interplay of the processes which control appendage and segment formation (Linder,
1952). This is confirmed in my data, most commonly with males and females from
the same sample ; the males tending to have a smaller number of appendages, the
same number of segments and so a higher number of apodous segments than the
females. Similarly, Lepidurus bilobatus has 33 segments, 60 pairs of appendages and
6 apodous segments (Linder, 1952), while L. batesoni with the same number of
segments, but only 39-52 appendages, has 8-9 apodous segments.
The apodous segments were frequently counted in early descriptions of Notostracan
species, and several specific distinctions have been based on small differences in this
number. It is now known that there is so much variation in this character in Triops
that it is useless as a systematic character ; the pure line T. cancriformis had a
variation of three (5-7). It may be valid in some Lepidurus, and is of use in at least
one specific distinction.
A few general rules can now be drawn from the data on segmentation and appendage
number.
In both genera males often have a higher number of segments within a sample,
while the reverse appears to be unknown. In 25 samples of Triops I found this to be
the case, while in 21 the difference was insignificant.
Certainly in Triops, and probably in Lepidurus, the males tend to have fewer
appendages and so a higher number of apodous segments than the females ; in 44
Triops samples the males had a higher number of apodous segments, in 7 there was
no difference and in 1 the female had more.
Specimens with high numbers of segments tend to have a high apodous number
in both genera.
Thus, it is obvious that these characters can be used in systematics only with a
great deal of caution, and appear to be of more use in Lepidurus than in Triops.
(4) Carapace
The shape and size of the carapace in Notostraca varies considerably and the
differences found have frequently been used in the past by S3 7 stematists to dis-
tinguish species. Ghigi (1921) used the carapace shape as one of his arguments in
separating Triops into two genera : Thriops and Proterothriops (sic) ; Barnard
(1929) considered the shape to be a distinguishing feature between South African
species of Triops, particularly between his Apus numidicus and A. namaquensis. On
A REVIEW OF THE NOTOSTRACA n
the other hand, Linder (1952) made no use of the carapace size and shape in his
revision of the North American forms. The carapace is one of the structures which
is usually described adequately in the earlier papers.
In each individual the growth of the carapace from the earliest larva is probably
isometric ; a number of individuals of Triops cancriformis and T. granarius were
measured during the whole of their growth period and it was found that the ratio of
5 IO
CARAPACE LENGTH , MM
3C
Fig. i. Growth coefficient (k) of females of Triops cancriformis from Hampshire — five
individuals.
carapace length to total length remained the same throughout growth. In the
former species the growth coefficient (k) of the carapace was 1/02, while in the latter
species k = roi ; these results are shown graphically in Text-fig. 1.
The basic variation in the carapace is its size relative to the total length of the
animal ; in some forms it covers a much greater length of the body than in others ;
correlated with this relative size-difference are other differences in shape and strength
12 A REVIEW OF THE NOTOSTRACA
of the carapace ; in those specimens in which it is relatively small it is always more
rounded in outline, flatter, and less strong than in those in which it is large. These
differences hold good for both genera.
The relationship between carapace length and total length depends both on
changes in size of the carapace — the number of segments that it covers — and on
changes in the number of abdominal segments. A comparison between two species
of Triops made on the living animals illustrates this ; British female T. cancriformis
have about 33 segments of which about 19 are exposed behind the carapace, which
thus covers some 14 segments ; T. granarius from Johannesburg have, in the female,
about 38 segments of which 27 are exposed, so that in this species the carapace
covers about n segments. In the second species the increase in the number of
exposed segments comes both from a shorter carapace and a larger number of
abdominal segments. It is impossible to make such calculations on preserved
specimens with any accuracy, because of contraction in the preservative.
The carapace appears to cover about 11-14 segments in most specimens, for on
these segments are borne the most anterior of the spines which occur on the margins
of the segments in the exposed portions of the abdomen ; in all the specimens
examined, the first of these spines appear on segments n-14, regardless of the total
number of segments present.
Within a species, males have smaller carapaces than females, although this
dimorphism is less marked in Lepidurus than Triops, and in shorter bodied than in
longer bodied specimens of the latter genus. This sexual dimorphism in the carapace
has been noted by many authors.
The dependence of the degree of this dimorphism on the number of segments
means that while it is usually possible to distinguish males from females at a glance
on this character within a population, males from short bodied populations may
actually have longer carapaces than females from populations with high segment
numbers. In the forms with the lowest numbers of segments the dimorphism may
be so slight as to be virtually non-existent.
The carapace is smaller, more round, and flatter in populations of both genera
which have relatively high segment numbers ; this is especially well marked in
Triops where the specimens with the highest numbers of segments have remarkably
small carapaces (Text-fig. 2).
Barnard (1929) maintained that the shape of the carapace was an absolute dif-
ference between his Apus numidicus and A. namaquensis , which he found in the
former species to be oval in shape and in the latter almost round ; he also gave data
on the number of apodous segments which show that A. namaquensis is longer in
the body than the other species, although some overlap occurred between the two.
He was concerned only with South African material, but on examining specimens of
these species — and of synonymous ones — from the whole of their range in Africa
and Asia I find that there is no discontinuity in the variation of carapace-shape.
The round carapaces and the oval are connected by populations of intermediate
form (Text-fig. 2A-e). The differences that Barnard found were differences between
the long and the short bodied forms of the same species.
I can find no differences in the carapace shape or size which are not correlated
A REVIEW OF THE NOTOSTRACA
i3
Fig. 2 Correlation of form with number of segments, a-e, Triops granarius, progres-
sive increase of segments (32-42) ; f, squared sulcus on some specimens of T. granarius
from South Africa ; g-h, T. longicaudatus , few and many segments respectively ;
1, Lepidurus apus ; j, L. batesoni. (3, 4, 5, — endites of first thoracic appendage,
cp = carapace, t = telson, fc = furca, sap — supra-anal plate).
i 4 A REVIEW OF THE NOTOSTRACA
with sex or the number of segments ; Triops cancriformis tends to have a lower
number of segments than T. granarius and so its carapace tends to be longer, less
flat, than in the other species ; and specimens of T. granarius with a low number of
segments have carapaces similar in shape to those of T. cancriformis. Lepidurus spp.
tend to have even shorter bodies and here the carapace may cover most of the
abdomen, and may be so deep as to enclose the animal laterally.
The shape of the sulcus, or posterior emargination, of the carapace has frequently
been described and importance attached to it ; I find it to be very variable and can
see no correlation with other characters ; it may be shallow and wide in both long
or short bodied forms, or small and round in similar animals. A peculiar squared
sulcus has been seen in several short bodied populations of Triops granarius from
Africa (Text-fig. 2f), but this grades into more rounded forms and is obviously of
no significance. Wide, shallow sulcus shapes occur most frequently in T. longicaudatus ,
but this is of very doubtful value in the systematics.
The difficulty of an accurate classification of sulcus shapes makes it unlikely that
this will ever be a useful character.
(5) Carapace armature
The carapace bears an armature of spines both scattered and localized. The whole
outside surface may be smooth, or may bear scattered upright spines ; the dorsal
carina frequently ends in a spine and may bear smaller spines along its length ; the
sulcus generally bears a row of marginal spines, and the outside edge of the carapace
may bear a similar row.
The carapace armature shows a great deal of variation, and has been referred to
very frequently in past descriptions ; some of this variation does seem to be of use
systematically, but it must be used with caution and only as a confirmatory character
in most cases.
The armature as a whole varies in its strength and development even within a
population, and in some animals the whole armature is more strongly developed
than in others — in the former not only are the spines larger and stronger, but they
are also more numerous. In Triops granarius, when the scattered surface spines are
well developed, the sulcus spines are particularly strong, the carina is denticulate
and the whole carapace is more rigid than in other specimens. In a population of T.
cancriformis from Tunisia (MNHNP) the armature of the exoskeleton is extra-
ordinarily weakly developed, though typical of this species in arrangement, and
extreme examples in this sample have no carinal or sulcal spines at all — a most
singular condition.
The scattered spines on the surface of the carapace were features used by Sars in
the description of two species of Triops (Apus trachyaspis and A. sculleyi, Sars,
1899), but Barnard found that specimens bearing such spines occurred sporadically
among South African material. I have found such specimens in samples of Triops
granarius and T. longicaudatus, both as isolated individuals and as complete samples.
Specimens in which this character is well developed are so conspicuous — the carapace
having a prickly feel — that I shall refer to them as the trachy aspis-i orm of whichever
species is involved.
A REVIEW OF THE NOTOSTRACA 15
The terminal spine of the carina is most prominent in larval Lepidurus] I have seen
specimens of L. arcticus, L. apus apus and L. apus viridis in which it is relatively
enormous in the second and third instars (Text-fig. 13A). In larvae of Triops it
develops later and is never as large as in young Lepidurus. The growth rate of this
spine in Lepidurus must be strongly negatively allometric, for although it is present
in most adults, it is relatively very much smaller in these than in the larvae ; in
Triops, when it occurs, this spine has a positively allometric rate for it is quite small
when it first appears, and becomes, in the adult, of a size relatively similar to that
of adult Lepidurus.
This terminal spine occurs in almost all adult Lepidurus and is absent in only a
few. There appears to be no correlation between its absence and other characters ;
L. batesoni and some specimens of L. apus apus and of L. apus lubbocki are without
it, but these specimens of L. apus are otherwise quite normal. There is probably a
variation in the growth rate of this spine to account for its disappearance in adults
of a few populations.
Fig. 3. Carapace of Lepidurus lynchi. a, typical form of carinal spines ; b, specimen
with no carinal spines.
In Triops there is a loose correlation between the occurrence of the terminal spine
and other characters ; within a species it is more commonly present in short than
in long bodied forms. There is some difference between species, too, and as might be
expected it is almost always present in the relatively short bodied species, T. cancri-
fortnis. It is present in the remaining species only in their short bodied forms.
The carina may bear a series of spines, most commonly at the posterior end just
anterior to the terminal spine. These have frequently been used in the past for
systematics (Ghigi, 192 1 ; Linder, 1952) and they have proved to have value in
certain cases. In Triops they may be present, or absent, in both long and short
bodied forms, but there are some interesting differences within T. cancriformis. In
this species a few small spines are generally present posteriorly, and these are more
numerous and much stronger in specimens from Morocco and Southern Spain ; these
are the populations referred to by Ghigi (192 1) as Thriops mauretanicus , but they
are now considered to comprise a sub-species (sens. Mayr, et al) of the more wide-
spread Triops cancriformis. Another sub-species of this species is characterized by
the complete absence of carinal spines.
In Triops australiensis and T. longicaudatus these spines are most frequently
16 A REVIEW OF THE NOTOSTRACA
completely absent, but in a number of populations, that from the rice fields of Biggs
County in California, for example, the whole carina bears an even row of very small
denticles.
Most specimens of Triops granarius are without carinal spines, and only a few of
the short bodied specimens have an arrangement like that of T. cancriformis. Very
rarely this species has a long row of very small denticles like those of T. longicaudatus.
The majority of specimens of Lepidurus have perfectly smooth carinae but a
remarkable series of large teeth along the carina occurs in L. lynchi, quite unlike
anything else in the Notostraca (Text-fig. 3).
The sulcus normally bears a marginal row of spines, the only exceptions being
some of the specimens of Triops cancriformis from Tunisia mentioned earlier. There is
a fairly clear correlation of the form of these spines with the number of segments in
Triops, but not in Lepidurus ; they are larger and fewer in number in short bodied
forms of the former genus.
Their development in Triops is fairly clear ; the spines at the outside angles of
the sulcus appear first, in the 4th or 5th instar, at which time the rest of the sulcus
has a finely granulated margin. The first marginal spines appear in the 7th or 8th
instar, and increase in number until about the 10th instar, after which time the
number is fixed, though in a few specimens small subsequent additions may occur.
There is little variation in this character in an individual after it is about 5*0 mm.
in carapace length.
Barnard (1929) considered that there was so much variation between individuals
in the sulcus spines that their use in systematics was not justified, although earlier
writers had placed much emphasis on them. Linder admitted this variation, but
thought that in some cases specific differences could be found.
I can discover no differences of the sulcal spines in Triops which can have any
value in systematics, most of them are correlated merely with body length ; in T.
granarius, short bodied forms have long, slightly curving spines and in longer bodied
animals a larger number of small, blunt spines occurs. As in other characters T.
cancriformis here resembles the shorter bodied T. granarius specimens.
In Lepidurus, a correlation with body length was not observed ; some specimens
with 28 and some with 33 segments had small squat spines, but the vast majority of
samples of all body lengths have long spines similar to those of Triops cancriformis.
An unusual arrangement occurs in Lepidurus apus packardi in which the margin
is closely set with many small squat spines like those of larval Triops.
The outer margin of the carapace in both genera normally bears a series of denticles
which produce a finely serrated edge ; this is variable in development, and is normally
stronger near the posterior angles of the carapace ; in only one case are these
marginal denticles of any value systematically ; some specimens of Lepidurus
lynchi bear a series of teeth along this edge which are very much larger than those
of any other known form.
The amount of individual variation which may be expected in the carapace
armature of a population was illustrated by the pure line of Triops cancriformis ;
here the number of posterior teeth on the carina varied from 2-10, the number of
sulcus spines from 24-32, and all the specimens had a large terminal carina spine.
A REVIEW OF THE NOTOSTRACA
17
(6) Supra-antennal crest
On either side of the ventral surface of the head there is a ridge, behind which is
set the first antenna ; this is the supra-antennal crest of Simon (1886), which Linder
(1952) suggests may be worth study. I find it variable within a single species, in
some specimens of Triops c. cancriformis it is denticulate and in others smooth and
less prominent. It seems improbable that it is of importance systematically.
(7) Eyes and dorsal organ
In all adult Notostraca the dorsal surface of the head bears a pair of compound
eyes, an ocellus, and the dorsal (or nuchal) organ.
EYE.
Fig. 4. Eyes and dorsal organs, a, Triops cancriformis ; b, T. australiensis
Triops granarius ; d, Lepidurus batesoni ; B,F,L.apus; g, L. arcticus.
c, c.
The size and arrangement of these structures are given in most specific descriptions
and considerable importance has been ascribed to them, in particular to the dorsal
organ ; Barnard (1929) showed that the dorsal organ varied in South African
material from a small round shape to a larger and triangular one. Further, he found
that the small, round dorsal organs tended to be set on a tubercle in his animals,
but that the triangular ones were less elevated and nearly flush with the surface of
the head. These differences, to which he ascribed much importance in separating
species, now appear to be another example of a character which is correlated with
the number of segments, and which varies in a similar fashion in several species.
In all the species of Triops the specimens with a low number of segments tend to
have small, round and elevated dorsal organs, and as the number of segments
increases so the dorsal organ becomes larger, less elevated, and more triangular in
shape (Text-fig. 4C-c 2 ).
This is well shown by Triops granarius ; in specimens from South Africa with only
33 segments the dorsal organ is small and round, or slightly pear-shaped, as it is in
zool. 3, 1. 2
i8 A REVIEW OF THE NOTOSTRACA
specimens with 36-37 segments from Chufoo, China (both BMNH) ; in longer
bodied specimens it is much larger and more triangular. In T. cancriformis the usual
shape is round, but in some of the longer bodied individuals it approaches the
triangular form of most specimens of T. granarius. T. longicaudatus is similar in
this character to T. granarius, but in T. australiensis a peculiar wide shape, with a
slight emargination of the posterior margin, occurs in a few specimens.
I have seen no specimens, and can find no records, of Lepidurus with triangular
dorsal organs ; here the round shape seems to be usual but in most species it varies
from round to oval (Text-fig. 4). In L. arcticus a peculiarly long, narrow oval shape
is common and this does not appear to occur in other species (Text-fig. 4G).
The growth rate of the dorsal organ is very strongly negatively allometric and
this further complicates its use as a systematic character. In the larvae it is a
relatively enormous structure so that in the first instar its median length is com-
monly about half that of the carapace rudiment (Text-fig. 13B) .
During growth to adult size its linear increase is only x 2 or x 3 while the relative
growth of the carapace is naturally very much greater. This negative allometry
appears to continue throughout growth.
Linder made use of the relative arrangement of the eyes and the dorsal organ in
his species of Lepidurus. He found that L. lynchi has the dorsal organ placed well
behind the posterior boundary of the eyes and of the tubercles over the eyes, in
contradistinction to the rest of the genus in which he found that the dorsal organ
was placed in part between the eyes. L. batesoni sp. n. has the first arrangement
(Text-fig 4D), but L. bilobatus — the other member of the long bodied group — has not.
In Triops, almost all the specimens examined had the anterior margin of the
dorsal organ between the eyes, the only exceptions being 8 of the 12 samples
of T. australiensis examined, in which the arrangement was precisely similar
to that of Lepidurus lynchi and L. batesoni (Text-fig. 4B) ; in the other four
samples it was normal. This may indicate that not too much reliance should be
placed on this character in either genus as a primary distinction between species.
(8) Telson
The telson bears an armature of spines on both dorsal and ventral surfaces, and
variation of these have commonly been used in the past for separating species
(Packard, 1883 ; Ghigi, 192 1; et at.).
The important spines on the dorsal surface of the telson fall naturally into four
groups to which it is convenient to apply names (Text-fig. 5) ; around the bases of
the furca are rings of f ureal spines ; on the posterior margin of the telson of the
larvae the first spines to appear are large and are identifiable in the adult — the
posterior marginal spines ; the median area of the telson may bear a row of large
spines or scattered smaller ones — the median spines ; around the dorsal sensory
setae are rings or arcs of setal spines.
The development, but not the origin, of these spines differs radically between the
several species of Triops (Text-fig. 5), and the final arrangement is of the greatest
importance in the systematics.
All the specimens of Triops which were available were examined on a geographical
A REVIEW OF THE NOTOSTRACA
19
basis, and no account was taken of previous determinations of the specimens ; it
was found that there was a strong correlation between the spine pattern of the telson
and the geographical distribution of the animals, but none with the sex or the
number of segments.
Fig. 5. Development of telson armature in Triops. a, larval stage (instar 2) common to
all ; b, c, d, instar 5-6 ; b v c v t> v 3-4 mm. carapace length ; b 2 , c 2 d 2 , adults. B n , T.
cancriformis ; c n , T. granarius ; D n , T. longicaudatus (the small arrows indicate the
position of the posterior marginals in each case ; /
spines) .
furcal spines, m = median
Without exception the specimens from Europe and western Russia have a small
number of median spines arranged in an accurate row in the centre of the telson, the
furcal spines are few and large, and the posterior marginals small, thin and remaining
on the margin in the adult (Text-fig. 5A) ; this pattern occurs also in North Africa,
the Middle East and northern India. In Africa south of the Palaearctic Region all
20 A REVIEW OF THE NOTOSTRACA
the specimens have a larger number of small, relatively scattered medians in the
mid-dorsal region, small, numerous furcal spines, and small posterior marginals
(Text-figs. 5c, 6a). This pattern overlaps the European type in North Africa and
the Middle East without forming intermediates, and then spreads across Central
Asia to the Chinese coast. In North and South America the medians are similar to
those of the European form, but there are two large spines, one on either side, at
the posterior end of the median row, which represent the enormously enlarged and
forwardly migrated posterior marginals, a fact which was confirmed by a study of
the larval development (Text-fig. 5D). This form also occurs to the exclusion of
others in the West Indies, the Galapagos, Oahu, and Japan ; a derivative occurs in
New Caledonia (Text-fig. 16).
Fig. 6. Telsons of Triops. a, T. granarius (trachyaspis-iorm) ; a v T. granarius ; b,
T. australiensis ; c, T. cancriformis mauretanicus.
The Australian forms have a very few small scattered medians, as in the African
form, but they are so few in number as to be frequently absent (Text-fig. 6b) ; the
rest of the armature is as in the African specimens.
This gives a picture of four large groups which are to a great extent allopatric,
but forming where they overlap no intermediates. The descriptions in the literature
completely confirm this grouping and are too numerous to list with profit.
As a systematic character this pattern appears to be perfect, there are no known
intermediates, it is unlikely that it has any direct adaptive significance, and it is
very easily seen in the specimens.
It must be remembered, however, that although the pattern is stable the numbers
of spines which form it are very variable ; in the pure line Triops cancriformis there
A REVIEW OF THE NOTOSTRACA
were 1-4 medians, 2-4 furcals, and, once, small additional posterior marginals
barely distinguishable from the primary pair.
The number of these spines is fixed from a size of about 5 mm. carapace length,
but as Text-fig. 5 shows, their relative sizes forming the true adult patterns are not
stable until a little later, and allowance must be made for this (e.g. Apus mavliensis
Fig. 7. Supra-anal plates of Lepidurus. a, L. apus apus ; a v L. apus lubbocki ; b, L.
batesoni ; c, <j>, c v $, Lepidurus arcticus.
22 A REVIEW OF THE NOTOSTRACA
Tiwari appears to have a different telson pattern from the other Indian forms, but
this species was described on immature individuals in which the posterior marginals
were still relatively very large).
The spines encircling the dorsal setae are very similar in arrangement, though not
in number, in all specimens of Triops examined, and no importance is attached to
them.
In Lepidurus, the spines corresponding to these four groups can be recognized,
but they are to some extent modified ; the posterior marginals are the first to appear
in the larval telson (L. apus) and the posterior margin of the telson on which they
are borne expands rapidly to form the supra-anal plate, carrying them with it, and
they are soon indistinguishable from the other spines on the margin of the plate in
most specimens. The f ureal spines are similar to those of Triops granarius and do
not differ much from species to species ; the setal spines are similar in form and
lack of variation to those of Triops ; the medians form an elongate row along the
mid-dorsal line of the plate.
This median series shows considerable variation and has frequently been used in
the separation of species ; the number of spines increases to a certain extent as the
animal grows (Linder, 1952) ; and this process probably continues until the animal
is at least 15-20 mm. in carapace length, to judge from a sample from Berlin of
Lepidurus apus (BMNH) consisting of large and small individuals.
The median spines may be borne on a slight keel, which is better marked in those
specimens with a large number of spines in this series ; it is impossible to draw a
line between presence and absence of a keel and contrary to Linder's opinion, I can
make no use of it.
The number of spines in the median series is useful in the systematics of Lepidurus
species, serving to distinguish the nominate race of L. apus from the other three
sub-species. L. arcticus has a much lower number than the rest of the genus. These
differences are not connected with the relative size in the adults of the supra-anal
plate ; both L. arcticus and L. lynchi have relatively low numbers of medians,
although the former has the smallest supra-anal plate in the genus and the latter
one of the largest.
The marginal spines of the supra-anal plate vary in size and number and there is
a connection between them and the median spines, when the latter are small and
numerous, so are the former ; the marginals therefore are relatively large and sparse
in Lepidurus arcticus, L. lynchi, and L. batesoni compared with the other species.
The size of the supra-anal plate itself varies between species — the most obvious
difference being in Lepidurus arcticus where it is very small — but a great deal of
the observed differences are due to age and sex ; males (Text-fig. 7c) have relatively
longer and more spatulate supra-anal plates than females (L. apus, Braem 1893 ;
L. arcticus, Somme 1934) ; and the structure has a positively allometric growth rate
throughout the period of growth (Braem, 1893 ; Campan, 1929).
The end of the plate is occasionally incised medianly, giving it a bilobed appearance ;
this appears in several species and has no importance (Linder, 1952), and it is to be
expected that this will occur in species other than those in which it has already been
recorded.
A REVIEW OF THE NOTOSTRACA
23
The shape of the telson was used by Packard (1883), but Linder has shown
that this is an unreliable character in Triops, though he records relative differences
in length and breadth in some Lepidurus species. The variation is such that no
reliance is placed on it here.
The dorsal sensory setae are present, and similar in form, in all species of Notostraca
that I have examined, and are longer in small than in large specimens.
(9) Segmental armature
Each segment which is exposed behind the carapace bears a series of spines on
its posterior border ; this series is interrupted by the appendages and is continued
vent rally only on the apodous segments.
The form of these spines on the ventral surface of the apodous segments has
frequently been used in the systematics of both genera ; Ghigi (1921) considered
them to be important in Triops, as did Linder (1952) in Lepidurus.
B
Fig. 8. Armature of apodous segments (a, b) and furca (c, d) of Triops. a, T. cancri-
formis without supernumeraries ; b, T. granarius with supernumeraries (s) ; c, $ T.
cancriformis or granarius ; d, <$, T. granarius.
On the ventral surface of these segments the spines near the midline are usually
smaller and set more closely together than the lateral ones. Linder (1952) considers
that the number and relative size of these centrals is diagnostic of certain species ;
he gives a variation of 13-28 teeth in Lepidurus packardi, L. couessi, and L. bilobatus,
and only 8 in L. lynchi ; I find the variation so great that this character is effectively
useless even though the differences are at times striking.
In Triops, the shape of the marginal spines was used by Ghigi (1921), who
found that they were squat in his T. mauretanicus, and finer in his other species ;
I find that the European and North African material — which Ghigi divided into
species — shows continuous variation in this character although, as he described, there
is a tendency for the Moroccan specimens to have squat spines.
These spines become rounded and heavy in males of Triops granarius in which
the ventral armature of furca and telson are heavy ; they are thus heavier in males
than in females of the same species.
A more useful character on the apodous segments is the presence or absence of
scattered supernumerary spines which occur between, and anterior to, the marginal
spines and are much smaller than them (Text-fig. 8). In Triops cancriformis only one
24
A REVIEW OF THE NOTOSTRACA
specimen has been seen which possesses any of these spines, a female from Palestine
(coll. Goldschmidt) which had 3-4 small supernumeraries on one of the apodous
segments. In the other species of Triops it is most unusual to find a specimen which
does not possess at least a few supernumeraries on each apodous segment, and in
the main there are 10-12 per segment.
I have not seen these spines in Lepidurus.
(10) Appendages
The appendages have been described in considerable detail for one species (Triops
cancriformis, Lankester 1881), but with the exception of one character have been
little used in systematics. The endites of the first thoracic appendage are drawn
1-5
o 1*0
0*5 -
5 IO 15
CARAPACE LENGTH, MM.
20
25
Fig. 9. Relative growth rates in Triops cancriformis from Hampshire. Furca and
longest endite of first thoracic appendage expressed as ratios furca/carapace and
endite /carapace for various carapace lengths. Any slope on the line thus indicates
allometric growth, since the growth rate of the carapace is isometric.
out into filaments of which the longest endite (the fifth) forms an antenna-like
structure in most forms. The length of the endite 5 and the number of segments of
which it consists have been described by too many past authors to list. Even
recently small differences in endite length have been considered to have significance
(Tiwari, 1952).
A study of the developing endite of Triops cancriformis and T. granarins has shown
A REVIEW OF THE NOTOSTRACA 25
that it grows positively allometrically and that this rate continues, to some extent,
throughout life (Text-fig. 9). So small differences in relative length are of no use
in systematics without much more investigation of the growth rates of this structure
in many populations.
It is likely that the length of the endites is not related to the size of the animal
per se, but to the instar that the animal has reached. Specimens which have grown
rapidly, with a large increment at each instar would have a different endite length
at a particular size from individuals with low instar increments due to poor condi-
tions ; this is borne out by the great size and relative proportional differences
seen in laboratory cultures.
In museum samples, perhaps for this reason, the individual differences in this
character appear to be greater than those to be expected between large and small
specimens ; if a sample is arranged in order of carapace-length the endite /carapace
ratio does not increase evenly up the series, as would be expected from the results
obtained from the growth of single specimens, and the inference to be drawn must
be that the specimens have grown at different rates and are of different sizes at
each instar.
This is further supported by the occurrence of two samples taken from the same
locality in different seasons ; two such samples from a locality in the Saone area of
France (MNHNP) collected two years apart consisted one of large, the other of
small adult individuals, but the endite /carapace ratio was very similar in each ; in
the first, individuals of 29-31 carapace length had a carapace /endite ratio of o*8o-
0*84 ; in the second sample the same ratio occurred in specimens of 9-5-10*8 mm.
in length.
The situation is further complicated by the fact that there is sexual dimorphism
in the endite length ; in 26 bisexual samples of Triops, 24 showed higher endite/
carapace ratio in the males. This may be due to the relatively shorter carapaces of
males than to any real difference in endite length relative to the total size of the
animals. In living specimens this appeared to be the case.
It is very noticeable that the endites are shorter relative to the body length in
both sexes of the longer bodied forms, but it is not possible to give reliable figures
for this because of the parallel differences in the relative carapace length which
renders inter-sample comparisons very difficult.
The range of the endite /carapace ratio is very great within a species determined
as such on other characters ; in Triops cancriformis this varies from 0*55-1*28, and
in T. granarius from 0*47-1*52. No correlation with the geographical distribution
could be found.
In Lepidurus the endites are not as long as in Triops. Lepidurus arcticus and L.
apus form a series from animals with very short endites which scarcely project
beyond the carapace, to those in which they are relatively long. In L. batesoni the
arrangement is unique ; the 5th endite of the first appendage is little longer than
that of the second appendage, and the 6th endite forms a claw in both appendages
although in all other Notostraca it is reduced to a small, soft lobe at the base of the
5th endite in the first thoracic appendage, undergoing negative allometry during
growth (Text-fig. 10).
26
A REVIEW OF THE NOTOSTRACA
Linder (1952) remarks "... the legs of various species are known to be very
similar to each other . . . ' ' ; I can find no reference to a comparative study of the
appendages of Notostraca, and so it seemed valuable to attempt at least a preliminary
survey. Unfortunately, it is necessary to dissect the appendages from the specimen
Fig. 10. Thoracic appendages : a, first thoracic appendage of Lepidurus arcticus ; b,
tip of second thoracic appendage of Triops cancriformis ; c, tip of first thoracic appen-
dage of T. cancriformis ; d, first thoracic appendage of Lepidurus batesoni ; e, endite 6
of first thoracic appendage of larval Triops cancriformis. (numbers = endites,
fl = flabellum, br = bract.)
in order to examine them properly, and this has naturally been possible with only
a few specimens. Appendages of representatives of both sexes of all four species of
Triops, and of several of Lepidurus were examined after mounting in polyvinyl
lactophenol.
A REVIEW OF THE NOTOSTRACA
27
The first antenna is present in all larvae and adults and is remarkably uniform
in structure ; it bears at its tip three setae in all species that I have examined,
although this number may be apparently altered by breakage. An aberration was
seen in one specimen out of a sample of a trachyaspis form of Triops granarius from
South Africa in which a pair of additional strong spines were present on one
margin.
The second antenna must be present in all larvae, for in them it is the main
locomotor organ ; the form and numbers of setae are identical in the larvae of all
four species of Triops, but I have been unable to make comparisons in Lepidurus ;
this appendage subsequently dwindles, due to negative allometry, and its locomotor
functions are taken over by the thoracic appendages. It is often absent in large
ttD
— ED
Fig. 11. Mouth parts of Triops cancriformis to show second maxilla, (md — mandible,
Mxi = first maxilla, Mxz = second maxilla, p = paragnath, T x = first thoracic
appendage, ED = efferent duct of shell-gland).
specimens, or it may be present (but very hard to detect) in the cleft in front of
the mandible. I have found both conditions in large individuals of all species and
can attach no importance to its presence or absence, contrary to the opinions of
Linder (1952) and Spencer and Hall (1896).
Ghigi (192 1 ) used the arrangement and relative sizes of the teeth on the triturating
surface of the mandible as a systematic character in his division of European and
North African material ; I have examined this character in many specimens con-
specific with his material and there is so much variation that the invalidity of the
character seems certain. Gauthier (1934) and Peres (1939) found similar variation
in their North African material.
The first maxilla has never been described comparatively ; I find the main varia-
tion to be in the row of spines on the ventral edge of the appendage. These tend to
be fewer in number and finer in Triops cancriformis than in the other species in the
28 A REVIEW OF THE NOTOSTRACA
genus, but there is probably a complete gradation of intermediates, and I can make
no use of the appendage in the systematics.
The second maxilla has received more attention than the previous appendage and
seems to provide an important character (Text-fig. n). Sars (1901) noted that
the second maxilla was larger in Lepidurus than in Triops ; this is so, and this
appendage is also larger in T. cancriformis than in the other species of the genus,
but even in this species is so small that it can scarcely be of importance in the feeding
mechanism. Linder (1952) reported that it was absent in large individuals of most
species of Triops {australiensis, granarius, numidicus, longicaudatus) , but that no
dwindling occurred during growth in T. cancriformis. I cannot agree with this. The
second maxilla is present in all samples that I have seen of T. cancriformis and T.
granarius, and absent in all those of T. longicaudatus and T. australiensis. Even in
very small specimens of the latter two species it is absent ; in a narcotized female
from Morawa in Australia it was clearly missing, yet this specimen was only 7-5 mm.
in total length.
In those species of Triops in which it is present it does not dwindle during growth ;
in a T. cancriformis of 8*o mm. carapace length the second maxilla was 0*2 mm. long
and in one of 18 mm. carapace length it was 0*4 mm. long ; in a T. granarius of
7*5 mm. carapace length it was 0*13 mm. long, and in one of 13*8 mm. it was 0*33 mm.
Thus, the presence or absence of a second maxilla provides a clear character
separating two species groups within Triops, but it is present and well developed in
all species of Lepidurus.
The efferent duct of the maxillary gland arises at the base of this appendage, or
from the position of the base if the appendage is absent ; this duct is longer in
Triops than in Lepidurus (Sars, 190 1) and longer in male than in female Triops, in
all the bisexual samples that I have examined.
The endites of the first thoracic appendage have already been dealt with, and I
could find no significant variation in the other lobes of this appendage, except
that the flabellum was relatively larger in Lepidurus batesoni than in other species —
another character of the 2nd thoracic appendage appearing on the 1st in this species.
In the second thoracic appendage, the form of the terminal " claw " (= exopodite,
Lankester 1881 ; apical lobe, Borradaile 1926 ; endopodite, Linder 1952 ; which
I shall call endite 6 from the precisely similar ontogeny in the early stages of this
and the other endites) was used by Ghigi (192 1) to differentiate between species of
Triops ; I find great variety in the form of this lobe, but none that can be correlated
with other characters. The relative lengths of endites 5 and 6 of this appendage
show a sexual dimorphism ; in males endite 5 is commonly longer than endite 6,
while in the females they are more nearly equal. This is well known and I have been
able to confirm it in all the bisexual samples that I have seen.
In the remainder of the thoracic series there is considerable variation in the shapes
of the lobes of the appendages, but I can find none that are of importance between
species ; however, all the specimens that I have seen of Triops granarius from
Central and Eastern Asia have endite 6 of the mid-thoracic appendages of a more
rounded shape than is usual.
The armature of these appendages shows much variation in numbers of spines
A REVIEW OF THE NOTOSTRACA 29
but not in their arrangement — a situation common in the armature on other
structures in these animals. The numbers of spines in any group tend to increase
during growth, but I have not been able to follow this closely.
The abdominal appendages showed no differences which might be of use systemati-
cally except to demonstrate once again a similarity between Triops cancriformis and
the Lepidurus species ; the flabellum of Lepidurus bears a number (20-30) of setae
around its outer margin, and these are reduced in number in Triops. But T. cancri-
formis has a higher number (10-20) than the other species (e.g. T. granarius, 4-9).
Linder (1952) mentions an inflated condition of the flabella in some specimens ;
this I find to be due to post-mortem changes, especially in those animals which have
died just after moulting, when the flabella are commonly so turgid with fluid after a
few hours that they have the appearance of small red balloons.
(n) Furca
The length of the furca has been considered an important systematic character
and was often included in even the early descriptions, but recent authors have thought
it unreliable (Linder, 1952).
The furca grow very rapidly in the larval stages of Triops but their positive
allometry soon becomes less marked, though some relative increase probably occurs
throughout life (Text-fig. 9). The furca are generally relatively longer in those
specimens in which the endites of the first thoracic appendage are long, and are
probably correlated in their development with the number of segments in the same
way as the endites. That there is no direct correlation with the endite length is
shown by Lepidurus arcticus in which the endites are very short, but the furca are
similar in length to those of the other species.
Gurney (1924) showed that sexual dimorphism in the armature of the furca
occurred in Triops ; the spines on the ventral surface of the proximal region of the
furca tend to be broader in males than in females, in extreme cases forming pro-
tuberant scales (Text-fig. 8c, d). With few exceptions this dimorphism is more
marked in long bodied forms, where the base of the furca tends to be relatively thick
and rapidly tapering. In Lepidurus these spines are a little, but not much, thicker
in males than in females.
PROTEIN SPECIFICITY
Oxyhaemoglobin has a characteristic absorption spectrum, and small differences
in the wave-length at which the axes of the absorption bands occur have been
demonstrated for several species of Daphnia by Fox (1945, 1946) ; similar
differences in the position of the absorption band axes of the chlorocruorin of species
and varieties of Sabella have been used systematically by the same author (1946).
The blood of Notostraca contains a considerable concentration of haemoglobin
in solution and the size of the animals is such that a volume of blood sufficient for
spectroscopic analysis can readily be withdrawn by a micro-pipette. It seemed
profitable to determine the wave-lengths of the absorption bands of as many
populations of Notostraca as were available, and to apply the results to the
systematics.
30
A REVIEW OF THE NOTOSTRACA
A Hartridge reversion spectroscope was used to determine these wavelengths, in
the manner described by Fox (1945), with a small refinement in technique to elimin-
ate personal bias and errors due to parallax ; the operator kept his eye to the instru-
ment throughout a series of measurements and an assistant made the readings on
the micrometer head and noted them down.
The wave-length of the axis of the oxyhaemoglobin-a band was determined by
comparison with a sample of blood of similar optical density and of known wave-
length (human blood, 5775 A). A number of readings were made on each sample of
blood and the results were treated statistically.
Several cultures of Triops cancriformis from different European localities were
compared, using several adults from each culture. The results (Table I) show that
although there is little individual variation, the three populations are apparently
different from each other; these differences gave a probability of significance of
0*02 when a t test was applied to them, and it may be taken that they are real and
not referable to errors in the methods of measuring.
Table I. — Wave-length of the axis of the oxy haemoglobin- cl band. Means of the readings
taken on each animal, the averages of these means (Xj, the number of readings
per sample(N), and the standard errors of the means ( ' S.E.J. The figures in
parentheses indicate the distribution of the readings within a sample.*
T. granarius
T. longi-
TV
</ m \'t*tP /•/i*i/ , -vi fnv*
>
m ij n n til c
J. fv\jyo lslA/rbl/r bj \jr rri>vo
A
Johannes-
Grahams-
6 il LtLLLll fA>5
Cali-
f
•\
Sweden
England
Italy
burg
town
fornia
Means per ani-
5779-7 (30)
5778-3 (40)
5777 -o (20)
5776-7 (10)
5778-2 (30)
5777-3 (30)
mal (A)
5778-9 (30)
5778-8(3o)
5778-2 (30)
5776-4 (30)
5775-5 (30)
5776-3 (30)
5779 -o (30)
5778-6 (20)
5777-9 (30)
5776-8 (30)
5776-5 (30)
—
5780-6 (30)
5778-8(32)
5777-3 (30)
5777*3 (30)
5778-3 (30)
—
5779-8 (28)
5778-9 (39)
—
5776-9 (30)
—
x(A) .
5779-6
5778-6
5777-6
5776-5
5777-1
5776-8
N
148
161
no
130
120
60
S.E.
0-356
0-305
0-281
0340
0-370
0-255
This indicated that inter-specific differences must be considerably greater than
these infra-specific ones to be of any value in systematics, and that many populations
of a species must be tested before a value characteristic of that species could be deter-
mined.
In fact, subsequent inter-specific comparisons gave results (Table I) very little
different from those obtained with Triops cancriformis.
The span between the axes of the oxyhaemoglobin and the carboxyhaemo-
globin-a bands was also examined, for this is known to show differences between
species in some cases (Fox, 1946). The results obtained with three species of Triops
showed no differences which would justify further investigation, (span : T. cancri-
formis = 50*1 A, T. longicaudatus = 48*2 A, T. granarius = 47*5 A).
The results of both these investigations are of no practical value in the systematics
of these animals — the differences being much smaller than those found and used in
Sabella by Fox — but do serve to demonstrate that in this character, as in others, the
species of Triops form a remarkably closely related group.
* A single specimen of T. australiensis from Kalgoorlie was tested and gave a result of 5779*3 A.
A REVIEW OF THE NOTOSTRACA
3i
REPRODUCTION
It has long been known that some populations of Notostraca contain both males
and females, and some only females. The occurrence of large populations in which
no males could be found has generally been assumed to indicate parthenogenesis,
especially since it was also known that isolated females from such populations could
produce viable eggs.
Bernard (1889) found scattered testis lobes in the gonads of female Triops cancri-
formis, Lepidurus apus and L. arcticus, and reported these occurrences as cases of
hermaphroditism. Zograf (1906) found ovarian tetrads developing in the testis
Fig. 12 Testis lobe in ovotestis of Triops cancriformis . 5(x section, Carnoy/Feulgen.
(5 = sperms ; Sc = spermatocytes ; F.D.C. = follicle duct cells).
walls of male L. apus, which must surely be an aberration of no use to the animal,
for the oocytes subsequently degenerated. Spencer (1896) could find no testis lobes
in the ovaries of Triops australiensis.
Bernards' findings, largely ignored since his account, have been confirmed in the
present work during investigation of the cytology of Triops and Lepidurus, and have
been reported elsewhere (Longhurst, 1954). Hermaphroditism has been found in
three species — Triops cancriformis, T. longicaudatus and Lepidurus arcticus — and it
was found that females of the first two laid viable eggs in isolation and contained
ovotestes (Text-fig. 12) ; no L. arcticus reached adult size. Females of Triops
granarius and T. australiensis were unable to lay eggs in the absence of males, but
32
A REVIEW OF THE NOTOSTRACA
did so readily as soon as males were put into the tanks with them and pairing had
occurred ; this was presumably correlated with the fact that no testis lobes were
present in the ovaries of the females of these species.
There is no evidence to show that these latter two species of Triops are ever any-
thing but bisexual ; in all the material I have examined of these species the large
samples invariably contained both sexes, and there are no records in the literature
of females occurring in large numbers without males. In T. cancriformis there appears
to be a complex situation : in the southern parts of its range bisexuality is normal,
and a female from Algeria had no testis lobes ; but in the more northerly regions
males occur sporadically, often in very low proportions, so that some of the females
Fig. 13. Notostracan larvae, a = carina of instar 3 Lepidurus apus ; b, metanauplius
of Triops cancriformis ; c, Neonatus of Lepidurus arcticus. {do = dorsal organ.)
probably reproduce without fertilization. In the extreme north of the range
(Britain, Sweden) males are unknown and here females contain ovotestes. Unfor-
tunately I have not been able to examine the gonads of females from a population
with sporadic male occurrence. Mathias (1937) gives a review of the occurrences
of males of this species.
In Triops longicauddtus the geographical distribution of males is not so clear;
however, specimens from Californian rice fields where males are unknown contained
ovotestes, and Linder (1952) reports examining many specimens from the Galapagos
Islands without finding males. Ueno (1935) records this species in Japan and found
no males in 78 specimens. It may be that all the populations of this species from the
Pacific region are hermaphrodite, for the small samples that I have seen from Hawaii
(Oahu) and from New Caledonia contained no males.
A REVIEW OF THE NOTOSTRACA
33
Reproduction in Lepidnrus is less well explored ; it is known (above) that herma-
phrodite individuals occur in some species, but males of these are also known in some
populations although their geographical distribution is not clear. The three long
bodied species all appear to be bisexual.
The size of the egg in Triops is variable, being larger in long bodied forms irres-
pective of species ; with this is correlated a size difference in the mature follicles in
&
6 9
9
©■
»9
Fig. 14. Notostracan chromosomes, a-c, 2 spermatocyte telophase ; d-e, young
oocyte ; f, resting unfertilized ovum ; g, spermatocyte diakinesis. a, Triops longi-
caudatus ; b, T. australiensis ; c, d, e, T. granarius ; f, T. cancriformis ; g, Lepidurus
apus. (g is half the scale of the remainder and is redrawn from Goldschmidt, 1953.)
the ovary : T. cancriformis (33 segments), 0*28 mm. ; T. granarius (38 segments),
0*45-0*52 mm. This size difference is correlated with size differences in the resultant
eggs and larvae, the long bodied forms having the largest larvae. In Lepidurus the
differences may be even more marked and are diagnostic of species in at least one
case ; the larvae of L. arcticus hatch at a later stage of development than those of
the other species (Text-fig. 13), and the eggs are very much larger, the ovarian
ZOOL. 3, 1. 3
34
A REVIEW OF THE NOTOSTRACA
follicles are relatively few in number, large and elongated in form, and completely
fill the space above the gut.
CYTOLOGY
Moore (1893) was the first to examine the Notostraca cytologically, and from obser-
vations on the divisions of somatic cells he concluded that Triops cancriformis was
amitotic (zn = 1).
While the present work was in progress, Goldschmidt (1953) recorded chromo-
somes in the testes of Lepidurus sp. from Palestine, finding that aceto-orcein squashes
gave diakinesis stages where n = 6 (Text-fig. 14G). I have since determined her
material as L. apus lubbocki.
Using testis smears I found that two populations of Triops granarius from South
Africa showed the number n = 4 ; this was most clearly seen in the telophase stages
of the spermatocyte (Text-fig. 14c). This number was confirmed in females of the
same populations in which a haploid set of chromosomes was readily seen in the very
early oocytes (Text-fig. 14D, e).
Sections of the ovotestes of Triops cancriformis from the European cultures showed
n = 4 in the oocytes, most clearly in the late oocytes or ova, resting in the longi-
tudinal oviduct (Text-fig. 14E) ; squashes of the same gonads showed n = 4 in
secondary spermatocyte telophases in the testis lobes.
In both of the above species of Triops the diploid number, 2n = 8, was confirmed
by finding mitoses in cells of the expanding follicle walls, at the stage of increase in
size of the oocyte and nutritive cells. In T. longicaudatus from California the number
n = 4 was seen in early oocytes (Text-fig. 14A) and in spermatocyte telophases, but
it was not possible to confirm the diploid number.
A single male of Triops australiensis from Kalgoorlie was available and testis
smears were made from it which showed, in the spermatocyte telophase, a comple-
ment of n = 5 ; this number was counted in at least 16 nuclei. The 5 chromosomes
Table II. — Chromosome numbers of Notostraca
Haploid
A
number
Diploid number
Females
f
Males
Females
L. apus lubbocki .
6
(diakinesis)
—
—
T. c. cancriformis
4
4
8
(2 spermatocyte
telophase)
(resting oocyte)
(follicle duct
epithelium)
T. granarius
T. 1. longicaudatus
T. a. australiensis
4
(2 spermatocyte
telophase)
4
(2 spermatocyte
telophase)
5
(2 spermatocyte
telophase)
4
(young oocyte)
4
(young oocyte)
8
(follicle duct
epithelium)
A REVIEW OF THE NOTOSTRACA 35
in this haploid set are all sub-equal in length (Text-fig. 14B), and it seems probable
that this situation could have arisen by the fragmentation of one chromosome in a
set of four, such as the other species possess, in which one is nearly twice as long as
the other three.
My findings and those of Goldschmidt are summarized in Table II, and both agree
in ascribing a low number of chromosomes to the Notostraca so far examined.
Moore's account of amitosis may be true of the cells that he studied, but even
that is doubtful in view of my chromosome counts in epithelial cells ; his inference
of a single chromosome was obviously incorrect.
All the Triops material was fixed in Carnoy's or Susa's fluids and was stained with
the Feulgen reaction.
BIOLOGY
Very many accounts of the ecology of the Notostraca and of their appearance in
temporary pools have been published, and a clear picture of their biology can be
derived from the literature.
There appear to be almost no ecological differences between the species of Triops,
a slight difference between Triops and most species of Lepidurus, and a marked
divergence between L. arcticus and the rest of the Notostraca.
So far as is known, Triops occurs only in waters which dry out regularly, and the
eggs normally hatch in the field only after a period of dessication. The habitat
itself may vary a great deal in size from tiny rain pools and cart-ruts to large tempor-
ary lakes (Main, 1953) and the water may be fresh as in the Hampshire locality
(Hobson and Omer-Cooper, 1935), brackish as in the Scottish pools (Balfour-Browne,
1909), or saline as in the lakes on the Tibetan plateau (Schlagintweit, 1872) and in
Australia (Main, 1953).
The farming practice of rice fields makes these ideal situations for Triops, which
are sometimes present in enormous numbers and have been recorded as a rice field
pest from many parts of the world (they uproot the rice seedlings) .
Lepidurus may occur in temporary pools, but there seems to be a tendency for
most species to live in waters which dry out less regularly than the Triops pools.
Lundblad (1920) records that Lepidurus apus is a spring form which appears when the
temperature rises early in the year in ponds or ditches which have held water all the
winter : Linder (1952), on the other hand, says that he has seen the same species
in Sweden in temporary pools. L. apus viridis occurs in a pool near Christchurch,
New Zealand, which dries out in normal years, but is only very rarely found in neigh-
bouring permanent ponds (G. Parry, personal communication). Lepidurus spp.
regularly occur in temporary, often alkaline, pools in North America (J. Lynch, in
litt.) . L. Glauert has sent me a map showing all the records for Triops and Lepidurus
in Western Australia and it is clear that Lepidurus is restricted to the south-western
coastal belt where there is regular winter rain and Triops to the arid interior where
rainfall is, at the most, sparse.
Lepidurus arcticus is confined to the boreal and alpine regions of the Holarctic,
where it may occur in large lakes and form an important food of Salmonidae and
where the eggs could never be dessicated (Somme, 1934), it is also in pools which
3 6 A REVIEW OF THE NOTOSTRACA
normally do dry out each year (J. Mohr, in litt.). The only other records of Lepi-
durus occuring in lakes refer to L. lynchi (Linder, 1952).
It has been thought that there is a difference between the two genera in the con-
ditions necessary for the hatching of the eggs : Fritsch (1866) and Grasser (1933)
thought that Triops eggs could hatch only after dessication, while Brauer (1877)
believed that the eggs of Lepidurus were incapable of withstanding dessication.
Schaeffer (1756) and Kozubowsky (1857) showed that the eggs of Triops were some-
times capable of hatching without drying out. This has been confirmed in the present
work and it is now known that the eggs of at least three species of Triops can hatch
out, after an interval for development, in the water in which they were laid, or will
remain viable if dried out and will hatch when replaced in water. This accounts
for the report by Mathias (1937) that Triops has two types of eggs : the one drought
resistant and the other not.
Fox (1949) reported that the eggs of Lepidurus apus viridis were able to hatch
after drying, and I have confirmed this with another mud sample from the same
locality as his ; L. arcticus from Iceland were also hatched from dried eggs. The
ecology of Lepidurus outlined above indicates that the eggs of this genus must in
many cases be drought resistant.
The eggs of Triops probably do not often hatch out without prior dessication,
as the water must have a low osmotic pressure for this to occur, a condition unlikely
to be found on the bottom of a pool where the eggs are laid, but most likely when the
pool refills with water after rain and the eggs float to the surface.
SYSTEMATICS
From the analysis of the characters used in the systematics of the Notostraca
it is evident that many of the obvious differences between individuals must be
correlated with differences in the number of body segments (Text-fig. 2) ; in Triops
there is no discontinuity in the variation of this number, so that neither the number
itself nor characters correlated with it can validly be used in the separation of species.
In this genus, as the number of segments increases to give a long bodied form so
the carapace becomes smaller, rounder and flatter ; the number of legs remains
about the same and consequently the apodous number becomes higher ; the furca,
and the endites of the first thoracic appendage become relatively shorter ; the dorsal
organ becomes larger, less elevated, and more often triangular in outline ; the sulcus
spines become smaller and more numerous, and the terminal spine of the carina is
more frequently absent ; sexual dimorphism becomes more marked.
Similarly, certain characters are correlated with the sex of the specimen ; males
tend to have more segments than females and so the characters outlined in the previous
paragraph vary from male to female within a population ; characters other than
these also show sexual dimorphism : the efferent duct of the shell gland is longer
in males than in females ; endites 5 and 6 of the second thoracic appendage are more
nearly equal in length in females ; the nth thoracic appendage bears a brood pouch
in females ; the ventral armature of the furca of males is coarse, often forming scales
rather than spines, and with this is correlated the ventral armature of the telson
and apodous segments, which vary in the same way between males and females.
A REVIEW OF THE NOTOSTRACA 37
These two considerations then — segment number and sex — invalidate a number of
characters in the systematics of Triops. With those that remain there appear to
be good grounds for dividing the genus into four groups, each with a geographical
basis. The valid characters appear to be : the armature of the telson, the presence or
absence of the second maxilla, and the arrangement of the eyes and dorsal organ.
Of these the most important is the armature of the telson, which is diagnostic for
each group ; the other characters, together with a few less well-marked ones, confirm
this primary grouping.
Two groups are partially sympatric, and here there is no tendency to hybridize
and no intermediates have been found in the areas of overlap ; there is a record,
moreover, of two species — which correspond to two of these groups — living in the
same pools in several localities in Morocco (Peres, 1939). These two forms, therefore,
behave precisely as biological species (sensu Mayr, 1942) would be expected to do
when they become sympatric ; it seems justifiable to consider them, therefore, as
species and since the degree of morphological difference between all four groups is
similar to that found between these two, then all four may equally well be regarded
as species.
Each species, then, has a geographical basis and a clear cut character in the telson
armature, but contains populations which are remarkably different in general appear-
ance depending on the number of body segments in the specimens. What determines
the body length is completely obscure, for there appears to be a general tendency
for the longer bodied animals to occur in the warmer regions of a species range, but
there are many blatant exceptions to this generality ; Barnard's Apus ovamboensis
is a population of exceptionally short bodied Triops granarius from a very hot and
dry part of Africa.
It is likely that an experimental study of the effect of environmental factors on
morphology would throw some light on this problem, for perhaps the effects are seen
only after a considerable number of generations.
Several species show some indications of the presence of geographical races ; the
differences between these are much slighter than the specific differences, and their
distribution indicates that the geographical barriers between them are very slight.
Triops cancriformis will serve as an example ; the populations in Morocco and
southern Spain differ in several respects from the rest of the species, even though
in Spain there is no geographical barrier where the change occurs, and there is some
evidence that intermediates occur where the races meet, for in Seville and near Gib-
ralter the specimens are identical with the Moroccan ones and from Valencia they are
similar to the specimens from the rest of Europe, but in Ciudad Real — midway
between the two areas — the specimens are intermediate. A similar zone of inter-
mediates seems to occur in Spanish Morocco east of Ceuta.
Ghigi (1921) considered these races of Triops cancriformis to be species and des-
cribed them as such ; Colosi (1922) and Peres (1939) referred to Ghigi's species as
varieties of the single species T. cancriformis but Gauthier (1934) used a true
trinomial nomenclature and considered them to be sub-species or geographical races ;
I follow this nomenclature and believe them to be sub-species in the sense that
Huxley (1942) and Mayr (1942) used the term.
3 8 A REVIEW OF THE NOTOSTRACA
The arrangement of the species seems to follow different principles in Lepidurus
from that in Triops, and the following account owes very much to the work of Linder
(1952) who demonstrated the existence of two species-groups within the genus, each
group characterized by its segment numbers.
The variation in segment number in Lepidurus is not so great as in Triops, and
many characters show no correlation with this number in Lepidurus though the
correlation is obvious in Triops : the armature of carina and sulcus ; the endites of
the first thoracic appendage ; the number of apodous segments and the size and shape
of the dorsal organ. The segment number shows a marked discontinuity at about
30 ; the vast majority of specimens have less than this number, and those which
have more are aberrant in other respects.
Within the short bodied group there is relatively little differentiation, but Lepi-
durus arcticus can be separated at once, and is distinguished by its range, its habitat,
its relatively short supra-anal plate and endites. The remainder of the group appears
to comprise a single species, with a few rather ill-marked, but very widespread, sub-
species.
Linder (1952) showed that Lepidurus couesii was conspecific with specimens deter-
mined as L. macrourus Lilljeborg. I can find no differences between the specimens
that I have seen of either of these species and those of the European L. apus. The
ranges of variation in the segment number are similar, the size and armature of the
supra-anal plate is the same, and so is the armature of the carapace.
In California a form occurs which is admitted as a species by Linder, Lepidurus
packardi ; in these specimens there is usually a slightly higher number of segments
than in the normal L. couesii (= L. apus), the apodous number tends to be higher,
and the sulcus spines are very small, numerous and closely packed. In the Medi-
terranean region there occurs a form which diverges from typical L. apus in a similar
way except that the sulcus spines are normal. Both of these forms replace the typical
race over very restricted areas, but there appear to be no effective geographical
barriers separating the several ranges from one another, and for these considerations
I propose to consider these forms to be sub-species of the typical and widespread
L. apus.
The Australasian forms do not differ from typical Lepidurus apus as much as do
the above two sub-species, and many specimens would be indistinguishable if placed
in a sample from Europe. However, I have seen no specimens which have the high
number of central spines on the supra-anal plate which are very common in the
typical L. apus, so that there is a slight degree of morphological differentiation and
I propose to consider the Australasian form as another sub-species, L. apus
viridis. The South American forms which I know only from descriptions must
be considered as another sub-species of L. apus.
The rest of the genus comprises the few longer bodied specimens which are known.
All have coarse and sparse marginal spines on the supra-anal plate — a character
which they share with Lepidurus arcticus — and a low number of central spines on the
same structure ; specimens from three localities bear uniquely large spines on the
carina and the carapace margin and these are Linder's L. lynchi, typical form and
var. echinatus. A small sample from Russia has a considerably higher apodous
A REVIEW OF THE NOTOSTRACA
39
number than the rest of this group and shares with L. lynchi a peculiar arrangement
of the eyes and the dorsal organ, which resembles the arrangement in Triops austra-
liensis where the anterior margin of the dorsal organ may be placed well behind the
posterior margin of the eyes. A single specimen from Utah was placed in this
species-group by Linder on account of its segment number and seems to agree with
the description of Lepidurus bilobatus Packard, having a normal carapace armature
and normal arrangement of the eyes and dorsal organ. The only possible arrange-
ment at the moment is to consider all these three forms as three separate species,
but it seems very probable that future work will be able to find a connection at least
between L. bilobatus and the Russian sample (L. batesoni sp. n.) and perhaps between
these and L. apus.
It will be noted from the foregoing account how widespread are the species of the
Notostraca and this is probably accounted for, as it is in some other invertebrates,
by their passive distribution ; the dried viable eggs must be blown around by wind,
and transport by birds is not unthinkable, for the eggs when laid are extremely sticky
and remain so for some days while the shell hardens, and so could presumably adhere
to larger animals, The eggs of other phyllopods are known to be capable of passing
unharmed through the guts of amphibia (Mathias, 1937), and birds are known to
eat Notostraca ; starlings (Decksbach, 1924) and gulls (Balfour-Browne, 1909)
are. recorded as feeding on Triops cancriformis, and Summerhayes and Elton (1923)
watched Arctic terns feeding Lepidurus arcticus to their young, and thought that
they might drop them accidentally into fresh pools on their way to the nest.
A passive distribution such as this must mean that geographical barriers are not
nearly so effective as they are for sedentary, or non-passively distributed animals,
and has produced species with world-wide distribution in other animals, such as
Tardigrades and Rotifers.
In addition, the group has a very long geological record and has had ample time
to occupy all suitable areas ; fossils from the Permian (Guthorl, 1934) are clearly
Notostracan carapaces, and forms from the Triassic of Europe (Trusheim, 1938)
are certainly Triops and differ from extant T. cancriformis only in the small size
of the terminal carinal spine. The upper Triassic L. stormbergensis (Barnard, 1929)
from South Africa is very similar to recent species except that the supra-anal plate
has, apparently, no central spines as in some extant L. arcticus.
It is impossible from these few fossils to give any account of the history of the
group, except to point out how little evolution has occurred in the space of 170
million years since the Triassic forms were alive.
However, something can be deduced about more recent changes in distribution.
During the Pleistocene glaciations Lepidurus arcticus was much more widespread
in western Europe than it is now ; this species is known from lacustrine beds of
that time from Scotland (Bennie, 1894) and from the Isle of Man (Geikie, 1894), and
now shows in Scandinavia the typical distribution of a boreo-alpine relict occurring
at sea level in the North and at progressively greater altitudes towards the South
(Somnie, 1934).
Triops cancriformis must have been absent from much of its present range at the
same period, and the post-glacial extension may have been performed largely by
4 o A REVIEW OF THE NOTOSTRACA
hermaphrodites, which predominate in the northern parts of its range ; for these
would be more efficient in dispersal as only a single egg would be required to effect
a colonization. Perhaps the increasingly sporadic occurrence of males towards the
north indicates a spread northwards of bisexuality.
All the known rice field populations consist entirely of females (or hermaphrodites
if parthenogenesis is assumed not to exist in these animals) and this strengthens
the above argument, for these are relatively new habitats for Notostraca, and a coloni-
zation by hermaphrodites ahead of the bisexuals seems to have occurred.
The westward extension of Triops longicaudatus across the Pacific from the largest
area of distribution in North America, may also be of relatively recent origin and
has apparently been performed again by hermaphrodites, for no males are known
from the Pacific populations (p. 47).
Nomenclature
Keilhack (1909) and Fox (1949) have shown that the generic name Apus Schaeffer,
1756 should be rejected in favour of Triops Schrank, 1803 ; this practice will both
accord strictly with the Rules of Nomenclature and will avoid further confusion in
this genus, and in the avian genus Apus Scopoli, 1777. I therefore propose to follow
Keilhack in the use of Triops.
The use of trinomials is necessary to describe formally the geographical sub-species
recognized.
The lists of synonymies before each species are not complete lists of references to
that species, for these would be too long and not very useful — but include (a) the
names, and the various spellings of each name, that have been applied to that species,
and (b) important descriptive works.
Apus has been applied to Crustacea other than Notostraca ; Apus pisciformis
Schaeffer is an Anostracan and Apus caudatus De Kay, was a crustacean parasitic
on a crab.
Identification
The following key should serve to identify specimens down to species, but the
user must bear in mind the extent of variation in any character and not expect the
figures to match the specimen in details of armature. It would be very valuable if
the segment number, the apodous number, and the number of appendages were
given in any future records.
It must also be remembered that every individual in a sub-species may not be
typical of it ; many ornithologists consider a sub-species to be valid if 75% of its
specimens can be placed in it without question. In Triops cancriformis , for example,
it may be difficult to ascribe single specimens to the nominate race or to T. cancri-
formis simplex, for the smooth carina which is typical of the latter occurs in some
individuals of the former, but I have no knowledge of whole samples of the nominate
race with smooth carinas.
Key to species of Notostraca
1. Supra-anal plate present (Text-fig. 7) . . . . . . . (Lepidurus) 2.
Supra-anal plate absent .......... (Triops) 6.
A REVIEW OF THE NOTOSTRACA 41
2. Segments more than 30 .......... . 3.
Segments less than 30 ........... 5.
3. Anterior margin of dorsal organ between eye tubercles (Text-fig. 4E) L. bilobatus (p. 53).
Anterior margin of dorsal organ well posterior to eye tubercles (Text-fig. 4D) . . 4.
4. Carina and/or lateral margins of carapace with large spines. (Text-fig. 3) Apodous
segments 3-5 . . . . . . . . . . L. lynchi (p. 53).
Without large spines on carina or lateral margin. Endite 6 of first thoracic appendage
as in Text-fig. iod. Apodous segments 8-9 .... L. batesoni (p. 54).
5. Supra-anal plate short, 0-5 spines centrally, marginals few (Text-fig. 7c). Endites
scarcely project beyond edge of the carapace . . . . . L. arcticus p. 52).
Supra-anal plate long (Text-fig. 7 a, b), 4-100 central spines, marginals numerous.
Longest endites with 3/4 of their length projecting beyond edge of carapace (Text-
fig. 2i) . . . . . . . . . . . L. apus (p. 50).
6. Second maxilla absent ........... 7.
Second maxilla present (Text-fig. 11). . . . . . . . . 8.
7. Posterior marginals sub-equal to medians and well forward of the margin (Text-fig.
5D 2 ). Medians large, 1-4 in number, in a row T. longicaudatus (p. 46).
Posterior marginals reduced and marginal. Medians small, scattered when more
than 3-4, often absent. (Text-fig. 6b) . . . . . T. australiensis (p. 48).
8. No supernumerary spines on the apodous segments (Text-fig. 8a) . Medians large, 1-4,
in a row (Text-fig. 5B 2 ) . . . . . . . . T. cancriformis (p. 41.)
Supernumerary spines present on apodous segments (Text-fig. 8b). Medians of
various sizes, scattered except when less than about 5 are present, when they form
an irregular row (Text-figs. 5C 2 , 6a, a x ) . . . . T. granarius (p. 44.)
Genus TRIOPS Schrank, 1803
1756. Apus Schaeffer (in part).
1758. Monoculus Linn, (in part).
1803. Triops Schrank.
192 1. Thriops (sic.) Ghigi.
1 92 1. Proterothriops Ghigi.
Triops is at once separable from Lepidurus by its lack of a supra-anal plate ; even
if a structure resembling a rudimentary plate (Text-fig. 7c) is found in some specimens
of Triops cancriformis (Linder, 1952) there is never any doubt as to which genus a
specimen belongs.
Proterothriops was erected by Ghigi for the reception of the long bodied forms of
the genus ; he was struck by the great difference in general appearance between
long and short bodied forms which I have now been able to show to be conspecific
in a number of cases.
Binoculus Geoffroy, 1762 and Apodis Zaddach, 1841, are not in a strict binomial
system and are ignored on this account.
Triopes Schrank, 1803, appears later in the publication than Triops and must be
regarded as a spelling lapse.
TRIOPS CANCRIFORMIS (Bosc)
1756. Apus cancriformis Schaeffer (in part),
1758. Monoculus apus Linn, (in part).
1801. Apus cancriformis Bosc.
42 A REVIEW OF THE NOTOSTRACA
1801
1803
1816
1871
1885
1885
1885
1909
1909
1909
1921
1921
1921
1922
1953
1953
Apus viridis Bosc.
Triops palustris Schrank.
A pus montagui Leach.
Apus himalayensis Packard.
Apus halicienis Fiszera.
Apus lublinensis Fiszera.
Apus varsovianus Fiszera.
Triops cancrifovmis (Bosc) Keilhack.
Apus cancrifovmis bidens Sidorov.
Apus carter if or mis transcaucasicus Sidorov.
Thriops simplex Ghigi.
Thriops mauretanicus Ghigi.
Thriops apulius Ghigi.
Thriops cancriformis var simplex, Colosi.
Triops cancriformis simplex (Ghigi, 1921) Margalef.
Triops cancriformis mauretanicus (Ghigi, 1921) Margalef.
Type. Original is unknown, and designation of neotypes desirable. A sample
from Kirkudbrightshire is selected (BMNH, 1907.10. 17. 1-4) and consists of a
number of neoparatypes.
Range. Western Europe (Spain to Sweden) east to Russia ; North Africa,
Balkans, Asia Minor, Middle East to India. Individual records too numerous to
list ; Lundblad (1920) gives many for Europe, Decksbach (1924) many for Asia.
Does not extend beyond 6o° N and range in Russia obscure, but no records authenti-
cated for Eastern Asia.
Habitat. Temporary fresh or brackish waters ; occurrence depends on the pools
filling when the temperature is high enough for development, so usually summer form
in Europe, spring form in N. Africa.
Rice field pest in N. Italy, Spain and La Carmargue.
Description. Head. — Dorsal organ round, oval or rarely triangular, small and
with its anterior margin between the eyes. Second maxilla present in all specimens
examined and relatively larger than in the rest of the genus.
Carapace. — Shape generally oval, more round in males. Carina with terminal
equal to, or longer than, sulcus spines (Text-fig. 15) ; sometimes number of smaller
spines on posterior carina. Sulcus spines long, 24-44, usually about 30.
Body. — Segments 32-35 in both sexes ; apodous 4-7 in ?£, 5-9 in <$<$. Apodous
segments without supernumerary spines on ventral surface.
Telson. — Median spines large, 1-4, in a single median row (Text-fig. 5b). Furcal
spines large, number sometimes varying from side to side of a single animal, and with
few scattered spines on lateral face of telson anteriorly.
Posterior marginals small, fine and marginal in the adult, and posterior margin
of telson sometimes drawn out to resemble a rudimentary supra-anal plate (Text-
fig. 6c).
Furca generally long.
Appendages. — 48-57 pairs recorded, but variation probably greater. Endites of
first thoracic appendage long.
Sexual dimorphism. — Not as well marked as in other species ; no males with furcal
scales seen. Abonyi (1926) records two sets of males in one summer in the same pool
A REVIEW OF THE NOTOSTRACA
43
of which the second batch had very reduced sexual dimorphism — so a dimorphism
of males may exist ?
Larvae. — Metanauplius (Text-fig. 13) ; dorsal organ of instar 1 generally round.
Reproduction. — Bisexual and hermaphrodite ; in the northern regions no males
occur and such females as have been examined are hermaphrodite. In the South
populations are bisexual and in central Europe males occur sporadically. Mathias
(1937) reviews occurrence of males.
Geographical Races
1. Triops cancriformis cancriformis (Bosc)
Range. Whole of species range with exception of that occupied by the other
two sub-species.
Description. Carina bears 0-10, generally 2-3, small teeth in front of the
terminal spine (Text-fig. 15E) and no large samples without specimens showing
these spines are known. Furcal spines small (Text-fig. 5B 2 ). Dorsal organ round-
oval.
Hermaphrodite and bisexual. Chromosome number zn = 8.
^Ssfe!
Fig. 15. Carinas of Triops. a-c, T. granarius ; d, T. cancriformis mauretanicus ; e,
Ej, T. c. cancriformis ; f, T. cancriformis simplex.
2. Triops cancriformis simplex Ghigi
Range. North Africa, Ceuta to Egypt.
Description. Characters of Thriops simplex Ghigi. Carina quite smooth in
front of the terminal spine (Text-fig. 15F) ; this is invariable in the specimens I
have seen from this area, and in those seen by Gauthier (1934), Colosi (1922) and
Ghigi (1921). Furcal spines small (Text-fig. 5B 2 ). These populations frequently
show a tendency to a general weakness in the strength of the armature and include
the sample from Kebili (p. 14) which has specimens with no carapace armature
whatever. The terminal spine of the carina is frequently reduced.
44 A REVIEW OF THE NOTOSTRACA
Apodous segments frequently higher in number than the nominate race ($? 5-7,
instead of 4-6).
Bisexual.
3. Triops cancriformis mauretanicus, Ghigi
Range. N.W. Africa : French Morocco and Tangier. S. Spain and Balearics
(Minorca).
Description. With the characters of Thriops mauretanicus Ghigi : armature
very strongly developed ; carina with a number of teeth posteriorly (Text-fig. 15D),
the largest often sub-equal to the terminal spine. Furcal spines very large (Text-
fig. 6c) ; ventral marginal spines on apodous segments very coarse. Apodous
number is similar to that of sub-species 2. Dorsal organ oval.
Bisexual.
Synonyms
Apus viridis Bosc was applied to figures of Schaeffer (1756) which showed juvenile
Triops cancriformis ; Triops palustris Schrank is presumably of this species because
of its N. European locality ; Apus montagui Leach, type in BMNH and clearly of
this species ; Apus himalayensis Packard, author's figures show clearly the cancri-
formis-type telson ; Apus varsovianus, etc. Fiszera, figures similarly show cancri-
formis-type telson ; Sidorov's two sub-species are insufficiently described and can
both be referred to this species ; Thriops apulius Ghigi ; this could be either sub-
species 1 or 2 as it was a single specimen with a smooth carina, but from its Italian
locality is probably of sub-species 1.
TRIOPS GRANARIUS (Lucas)
1864. Apus granarius Lucas.
1865. Apus numidicus Grube.
1877. ? Apus dispar Brauer.
1877. Apus sudanicus Brauer.
1880. Apus dukeanus Day.
1886. Apus bottegoi Prato.
1886. Apus namaquensis Richters.
1893. Apus sudanicus var. chinensis Braem.
1893. ? Apus sudanicus var. braueri Braem.
1893. Apus numidicus var. strauchii Braem.
1893. Apus numidicus var. dybowskii, Braem.
1895. Apus somalicus Wedenissow.
1899. Apus bottegoi, Bouvier.
1899. Apus trachyaspis Sars.
1899. Apus sculleyi Sars.
1899. Apus namaquensis Sars.
1899. Apus granarius Sars.
1907. Apus elongatus (nom. nov. for namaquensis Sars) Thiele.
1920. Apus zanoni Colosi.
1921. Apus asiaticus (nom. nov. for granarius Sars) Gurney.
A REVIEW OF THE NOTOSTEACA 45
1922. Triops uebensis (nom. nov. for bottegoi Bouvier) Co'osi .
1924. Apus ovamboensis Barnard.
1927. Apus numidicus var. sinensis Ueno.
1929. Apus cancriformis , Barnard, (non Schaeffer, 1756).
1929. Apus sudanicus, Barnard. (? Brauer, 1877.)
1934a. Apus granarius, Gauthier.
1937. Apus sudanicus, Gauthier. (non Brauer 1877).
1939- Apus numidicus, Peres.
1940. Apus sinensis Ueno.
1952. Apus mavliensis Tiwari.
1952. Apus orientalis Tiwari.
Type. MNHNP ; $ Holotype, unregistered other than label " Type A. granarius
Lucas ".
Range. South Africa to China ; the whole of Africa except for unsuitable forest
regions in west and centre ; Middle East, India, central and eastern Asia to the north
Chinese coast. Single sample in MNHNP labelled " near Paris " almost certainly
mistake in labelling.
Habitat. Temporary fresh and brackish waters ; unknown from rice fields ;
Peres (1939) compares its distribution in North Africa with that of Triops cancri-
formis and finds latter in steppe zones (300 mm. annual rain) and in the sub-steppe
(300-500 mm.) while present species is restricted to the sub-steppe.
Description. Shows great variation in body length and includes very short and
long bodied forms with all the associated variation.
Head. — Dorsal organ triangular, oval or round in the short bodied forms, anterior
margin between the eyes. Second maxilla present in all specimens examined except
in one case (see below).
Carapace. — Shape variable, round to oval. Carina with terminal spine in short
bodied forms only (Text-fig. 15), generally smooth. Sulcus round (or squared in
some African short bodied forms), spines many, often reduced in size, 24-72.
Body.— Segments : $$, 32-42 ; <?<?, 32-43- Apodous : ?? 4-13 ; £<£, 6-14.
Apodous always with varying number of supernumerary spines vent rally (Text-fig.
8b).
Telson. — (Text-fig. 5C 2 , 6a). Medians small, numerous, scattered, but when low
in number may form a rough row medianly, though this never as accurate as in
Triops cancriformis, and spines smaller. Furcals small and with many scattered
in front of them. Posterior marginals small, squat and marginal. Furca variable
with body length.
Appendages. — 44-46 recorded, but variation probably much greater than this.
Endites of first thoracic appendage variable with body length (Text-fig. 2).
Sexual dimorphism. — Strong in many long bodied specimens, males having strong
scales on furcal (Text-fig. 8d).
Larvae. — Metanauplius, those seen having in instar 1 a trapezoid-shaped dorsal
organ which later changes to triangular.
Reproduction. — Bisexual. Chromosome number 2n = 8.
46 a review of the notostraca
Geographical Races
With present knowledge not possible to recognize any with certainty. Some indi-
cations known — many South African (not S.W.A.) and Middle East (Bombay-
Baghdad) specimens have reduced sexual dimorphism even in long bodied forms ;
these correspond to Apus asiaticus Gurney and this name may eventually be applied
to a race with a restricted range. Some specimens from Eastern Asia have reduced
second maxilla, and this is even absent in few out of large sample (BMNH 1935.
6.18. 7-12) and these might be separated off as another race, but evidence too weak
at present.
Synonyms
Apus sudanicus Brauer belongs here doubtfully as the description is inadequate
and I have not seen the types, but Barnard's specimens determined as this are cer-
tainly Triops granarius. A number of species are placed here on evidence of original
descriptions : A. numidicus Grube, Braem's varieties of A. numidicus and A. sud-
anicus (exc. var. braueri which is doubtfully here for it refers to Brauer's description
of A. sudanicus), A. somalicus Wedenissow, Sars' three species (1899), A. granarius
Sars and the nom. nov. asiaticus Gurney, Ueno's var. sinensis and A. sinensis. I
have seen the types of A. granarius Lucas, A. bottegoi, Bouvier, Apus ovamboensis
(= cancriformis sens Barnard), and of Tiwari's two species. A. mavliensis Tiwari,
in spite of its odd appearance is clearly only a young form of this species. A.
bottegoi Prato is insufficiently described and may belong here or to the previous
species.
TRIOPS LONGICAUDATUS (LeConte)
1846. Apus longicaudatus LeConte.
1852. Apus domingensis Baird.
1 87 1. Apus aequalis Packard.
1871. Apus lucasanus Packard.
1871. Apus newberryi Packard.
1907. Apus frenzeli Thiele.
1916. Lepidurus patagonicus, Bruch. (non Berg, 1900).
1944. Triops pampaneus Ringuelet.
1947. Apus biggsi Rosenberg.
1947. Apus oryzaphagus Rosenberg.
1952. Apus longicaudatus, Linder.
Range. Western North America, south of 50 N, through Central to South
America, where only very scattered records (Thiele, 1907 ; Bruch, 1916 ; Ringuelet,
1944 ; Linder, 1952). W. Indies, Galapagos Islands, Hawaii, Japan and New Cale-
donia.
Habitat. Temporary fresh waters, rice fields in California and Japan.
Description. As variable as the preceding species, but not so many short bodied
forms seen.
Head. — Dorsal organ usually triangular, round in short bodied forms, and anterior
margin between the eyes. Second maxilla absent in all the specimens seen.
A REVIEW OF THE NOTOSTRACA 47
Carapace. — Text-fig. 2G., h). Shape varies from oval to round with segment
number. Terminal spine of carina very small when present, but generally
absent. Carina finely denticulate along whole length in some specimens. Sulcus
tends to be broad and shallow, spines variable in size, 24-60.
Body.— Segments : $?, 35-43 ; $S, 35-44- Apodous segments : $?, 5-12 ; <?<?,
10-13 in material examined. Apodous segments with varying number of super-
numerary spines on ventral surface.
Telson. — (Text-fig. 5 D 2 -) Medians large, 1-3, in a single row in the midline. Furcal
spines smaller than those of T. cancriformis. Posterior marginals very large, set
in the adult well forward of the margin, often pointing vertically. This arrangement
of marginals unique and all specimens of this species have it.
Appendages. — 54-66. (Linder's data mainly) : endites of first thoracic appendage
variable with segment number.
Sexual dimorphism. — Well marked in specimens with high segment numbers,
males then having furcal scales.
Larvae. — Metanauplius, indistinguishable from that of T. granarius.
Reproduction. — Bisexual and hermaphrodite, the latter only in California and
Pacific region populations.
Fig. 16. Telson of Triops longicaudatus intermedins, (pm = posterior marginals).
Geographical Races
1. Triops longicaudatus longicaudatus (LeConte)
Range. That of species with exception of New Caledonia.
Description. With the characters described above. There may be a difference
in the shape of the posterior margin of the telson on the forms from Galapagos and
Hawaii as against continental American forms ; in the former the posterior margin
is quite straight with no trace of an emargination in many specimens. This may
well be evidence for a sub-species inhabiting these islands. Chromosome number
2W = 8.
2. Triops longicaudatus intermedins subsp. n.
Range. New Caledonia. Two samples in MNHNP labelled " 93-1887 " and
48 A REVIEW OF THE NOTOSTRACA
" coll. Simon 25-96 " respectively are of this form and no others from the island are
known to me. These specimens, all females, form the paratypes of the new sub-
species. The name suggests that in their telson armature they are intermediate in
form between their nominate race and the rest of the genus.
Description. Second maxilla absent, segments 39, apodous 8-9. Dorsal organ
small, triangular to round. Carapace oval, terminal spine of carina very reduced,
sulcus spines long, 24-30. Few supernumerary spines on apodous segments ventrally.
Telson : medians large, 2-3, in mid-line, posterior marginals much smaller than in
nominate race but well forward of the margin (Text-fig. 16). This is the most char-
acteristic feature.
Synonyms
Apus obtusus James, 1823 and Apus guildingi Thompson, 1834 are clearly of this
species on geographical grounds, but neither description is detailed enough to confirm
this. Linder has shown that Packard's and Rosenberg's species are synonyms of
this species. The diagrams of A. frenzeli and Triops pampaneus both show the
longicaudatus-type telson, and Bruch's record of Lepidurus patagonicus is accom-
panied by a figure which certainly shows a Triops, and probably T. longicaudatus ,
though the telson is not very clear.
TRIOPS AUSTRALIENSIS (Spencer and Hall)
1896. Apus australiensis Spencer and Hall.
1905. Apus sakalavus Nobili.
1907. Apus madagassicus Thiele.
1 9 1 1 . Triops gracilis Wolf.
191 1. Triops strenuus Wolf.
Range. The drier regions of Australia, where it may be locally very common.
Madagascar.
Habitat. Temporary fresh water, sometimes (Wolf, 191 1 ; Main, 1953) in saline
or alkaline pools and lakes.
Description. Includes extremes of long and short bodied forms ; from Australia
several samples have been seen with aberrations (not damage) to limbs and lobes
of limbs ; both females in a sample from Ayer's Rock have reduced nth thoracic
appendages so that these are but a stump with a duct running through it on one side
of each specimen. Several samples from W. Australia have endites 5 and 6 reduced
to rudiments on one side of the thoracic series.
Head. — Dorsal organ triangular, ovoid or " wide ", often set with the anterior
margin well behind the eyes (Text-fig. 4B). Second maxilla absent in all specimens
seen.
Carapace. — Shape variable, terminal carinal spine usually absent and carina
frequently denticulate along the whole length. Sulcus round, often small, spines
variable in size, and numerous, 28-62.
Body.— Segments : ??, 35-43 ; $$, 36-44- Apodous: $$, 5-12; $$, 9-13.
Apodous segments with varying number of supernumerary spines ventrally.
A REVIEW OF THE NOTOSTRACA 49
Telson. — Medians small, scattered, fewer than in T. granarius, often very few or
even absent (Text-fig. 6b) ; furcal spines small, posterior marginals small, squat
and marginal in the adult. Furca varies with the segment number.
Appendages. — 48-66, but only few specimens counted. Endites variable with
segment number.
Sexual dimorphism. — As in previous two species.
Larvae. — Metanauplius, indistinguishable from that of T. granarius.
Reproduction. — Bisexual.
Geographical Races
1. Triops austr alien sis australiensis (Spencer and Hall)
Range. Continental Australia.
Description. With the characters described above. Chromosome number
2n = 10.
2. Triops australiensis sakalavus (Nobili)
Range. Madagascar.
Description. Very similar to the nominate race, but median spines are rather
more numerous in the specimens I have seen in MNHNP, which otherwise are clearly
of this species, and not of T. granarius ; the specimens have no second maxillae,
the carinae are denticulate, and the median spines are relatively sparse compared
with T. granarius.
Synonyms
Wolf's species are clearly Triops australiensis, showing the typical telson pattern
and Main has already (1953) suggested uniting them with it. T. madagassicus and
T. sakalavus are placed here on the assumption that only one species will occur in
Madagascar, since sympatric species are the exception in the Notostraca, but future
work will be needed to confirm this arrangement.
Genus LEPIDURUS Leach
1756. Apus Schaeffer (in part).
1758. Monoculus Linn, (in part).
1 8 19. Lepidurus Leach.
1924. Bilobus Sidorov.
Distinguished at once from Triops by the presence of a supra-anal plate. In
general, also, the segment number is lower, the carapace is longer and more com-
pressed laterally, the sexual dimorphism is weaker than in the other genus.
Bilobus Sidorov was erected on the erroneous assumption that median incision
of the posterior margin of the supra-anal plate was important systematically (see
p. 22, and Linder, 1952).
ZOOL. 3, 1. 4
5 o A REVIEW OF THE NOTOSTRACA
LEPIDURUS APUS (Linn.)
1756. Apus cancrifovmis Schaeffer (in part).
1758. Monoculus apus Linn, (in part).
1 801. Apus pro ductus Bosc.
1 81 9. Lepidurus productus Leach.
1850. Lepidurus viridis Baird.
1866. Lepidurus angasi Baird.
1873. Lepidurus lubbocki Brauer.
1875. Lepidurus couessii Packard.
1877. Lepidurus macrourus Lilljeborg.
1879. Lepidurus viridulus Tate.
1879. Lepidurus kirki Thompson.
1879. Lepidurus compressus Thompson.
1886. Lepidurus packardi Simon.
1893. Apus extensus Braem.
1900. Lepidurus patagonicus Berg.
1909. Lepidurus apus Keilhack.
191 1. Lepidurus hatcheri Ortmann.
1921. Lepidurus barcaeus Ghigi.
1952. Lepidurus couessii, Linder.
Range. Europe (excluding Britain), North Africa, Palestine, Asia Minor,
Russia ; North and South America ; New Zealand and Australia.
Habitat. Temporary fresh waters, but perhaps less restricted to those which
regularly dry out than, for instance, Triops cancriformis. Occurs in alkaline pools
in North America.
Description. This species has the largest range of any known Notostracan
and shows very little variation over the whole area.
Head. — Dorsal organ round or oval (Text-fig. 4E, f), anterior margin between the
eyes with the possible exception of the specimens seen by Ortmann (1911) from Pata-
gonia. Second maxilla present in all the specimens seen by me.
Carapace. — Oval, fairly compressed laterally but not so markedly as in next
species. Terminal spine of carina only rarely absent. Sulcus spines usually long,
rarely [packardi) reduced. Normally 35-50 in number.
Body. — Segments, 26-29 > Apodous, 4-6. No supernumerary spines on apodous
segments.
Telson. — Supra-anal plate relatively longer than in next species, the ratio carapace/
supra-anal plate being about 4-6 in adults. The marginal spines of the plate are
small and numerous in all except the South American forms. Central spines
4-100 or more, the higher numbers being borne on a distinct keel (Text-fig. 7 a) .
Appendages. — 35-48 ; endites of first thoracic appendages reach to the end of
the carapace in some, in others little longer than L. arcticus.
Sexual dimorphism. — Males occur, and the supra-anal plate is longer and more
spatulate in these ; otherwise the dimorphism is weak.
Larvae. — Metanauplius in the European and New Zealand forms and probably in
all. May differ from that of Triops by presence of rudiments of paired eyes in the
first instar (Bernard, 1892 : 158).
Reproduction. — Bisexual and hermaphrodite.
a review of the notostraca 51
Geographical Races
1. Lepidurus apus apus (Linn.)
Range. Europe (exc. range of next ssp.), Asia, North America (exc. California).
Description. Segments, 26-28 ; apodous, 4-5 ; supra-anal plate with 20-100
spines generally borne on a keel ; marginal spines of plate fine and numerous.
Bisexual and hermaphrodite.
2. Lepidurus apus lubbocki Brauer
Range. N. Africa, Palestine, Syria, Italy, Sicily.
Description. Segments, 27-29 ; apodous, 5-6 ; supra-anal plate (Text-fig. 7B)
with fewer central and marginal spines than above ssp (3-20 centrals) and keel less
prominent. Specimens from eastern part of the range tend to have the fewest
central spines ; endites relatively longer than nominate race. Bisexual. Chromo-
some number, 2n = 12 (Goldschmidt) .
3. Lepidurus apus packardi Simon
Range. California.
Description. Segments and supra-anal plate similar to sub-species 2. Sulcus
differs from that of all other forms by having very many small spines forming a
granulated margin. Bisexual.
4. Lepidurus apus patagonicus Berg
Range. South America (Chubut Territory, Patagonia).
Description. Segments, 29 ; apodous, 5. Marginal spines of supra-anal plate
coarser and fewer than previous forms, central spines few. Sulcus spines as typical
race. Bisexual.
5. Lepidurus apus viridis Baird
Range. New Zealand, Tasmania, and coastal or better watered regions of
Australia.
Description. — Very similar to nominate race. Segments, 27-28 ; apodous, 4-5.
Generally with low number of central spines on the plate (5-10), but one specimen
from Tasmania had more than 20 ; average is much lower than in nominate race,
however. I have seen no males.
Synonyms
I have examined the following types, and am satisfied that they are correctly
placed here : L. packardi, L. extensus, L. barcaeus, L. angasi, L. cotnpressus, L. kirki,
L. viridulus. The rest are included on the basis of the original descriptions or on
material previously determined.
Lepidurus lemmoni (Holmes, 1894). Holmes' description does not enable deter-
mination of his specimens with any here recognized, yet contains nothing to indicate
52 A REVIEW OF THE NOTOSTRACA
good differences from them ; the types are lost (Linder, 1952), and therefore it is
advisable to abandon the species.
LEPIDURUS ARCTICUS (Pallas)
1793. Monoculus arcticus Pallas.
1883. Lepidurus glacialis Packard.
1892. Lepidurus spitzbergensis Bernard.
1893. Apus productus var. glacialis Braem.
1896. Lepidurus glacialis, Sars.
1927. Lepidurus ussuriensis Sidorov.
Range. Circum-polar Arctic regions ; Aleutians, North America ; Alaska to
Labrador, Greenland, Iceland, Bear Island, Spitzbergen, Northern Palaearctic ;
Scandinavia to Siberia.
Habitat. Temporary fresh-water pools, the streams connecting pool systems,
and large lakes which are permanent. A reservoir in the Norwegian mountains
(Somme, 1934).
Description. Head. — Dorsal organ oval, sometimes very elongated, anterior
margin just between the eyes (Text-fig. 4G). Second maxilla present.
Carapace. — Oval, laterally very compressed. Terminal carina spine present, long.
Rest of carina smooth.
Body. — Segments, 26-28 ; apodous, 4-5. Apodous without supernumerary spines
on ventral surface.
Appendages. — 41-46, but variation probably greater. Endites of the first thoracic
appendage very short, scarcely reaching beyond margin of carapace (Text-fig. ioa).
Telson. — Supra-anal plate very small ; carapace /plate = about 12 in adults
of 20 mm. carapace length. Median spines very sparse (0-5) and marginals few
and coarse (Text-fig. 7c).
Sexual dimorphism. — Males rare, but known to have longer and more spatulate
supra-anal plates.
Larvae. — Post-metanauplius (Poulsen, 1940, and my cultures), equivalent to
about instar 3 of Triops larvae (Text-fig. 13).
Reproduction. — Bisexual and hermaphrodite. Males known from Bear Island
in very low proportion of the population. (Somme, 1934)-
Geographical Races
None could be recognized in the material available, nor could Linder (1952) find
any subdivisions of his material. Lepidurus ussuriensis Sidorov appears to be of this
species, for the endites are shorter than is usual in other species, and the supra-anal
plate is small (carapace /plate = about 9) and typical of this species in its armature.
However, the endites are projecting more beyond the carapace than is usual in
L. arcticus and may indicate a difference between this East Siberian form and the
typical Arctic forms. The weakness of the carina and the elongate form of the dorsal
organ confirm the placing of this species in L. arcticus.
A REVIEW OF THE NOTOSTRACA 53
LEPIDURUS BILOBATUS Packard
1883. Lepidurus bilobatus Packard.
1952. Lepidurus bilobatus, Linder.
Range. North America ; Utah, Colorado, probably Arizona (Linder, 1952).
Habitat. Not known, but occurs in arid areas so presumably in temporary
pools and lakes.
Description. Known only from Packard's description, and two further specimens
ascribed to the species by Linder.
Head. — Dorsal organ round or oval, anterior margin set between the eyes.
Carapace. — Arrangement of sulcus spines normal.
Body. — Segments, 33 ; apodous, 6.
Telson. — Supra-anal plate with numerous small marginals, 4-6 centrals.
Appendages. — 60 ; endites of first thoracic appendage as in Lepidurus apus.
Sexual dimorphism. — Males unknown.
Larvae, Reproduction. — Unknown.
LEPIDURUS LYNC HI Linder
1952. Lepidurus lynchi Linder.
1952. Lepidurus lynchi var. echinatus Linder.
Types. Holotype (?) and allotype ((J) in Uppsala Museum ; paratypes USNM
82101 ; var. echinatus. Holotype (?) USNM 82068, allotype ((J) USNM 82069.
Range. North America ; Washington, Nevada, and Oregon.
Habitat. Apparently in lakes (Linder, 1952 . . . North end of Goose Lake in
water two feet deep, muddy), but probably also in temporary pools.
Description. Long bodied form with unique carapace armature.
Head. — Dorsal organ round with the anterior margin set well behind the eye
tubercles.
Carapace (Text-fig. 3). — Oval. Carina absent except when it bears series of
unusually large spines. Specimens from first two localities have 0-20 of these carinal
spines. Some of those from Oregon have series of large spines along posterior part
of lateral margin in addition to carinal spines and carapace shape may be more round
in these. But no useful purpose is served by the formal term var. echinatus for these
latter forms ; L. lynchi is obviously characterized by the possession of large spines
on carina and margin and there is much variation in the distribution of these spines ;
even in the specimens from the first two localities the margin of the carapace bears
larger spines than is usual in Lepidurus.
Body. — Segments, 31-34 ; apodous, 3-5.
Telson. — Supra-anal plate about same size as in L. apus, central spines 2-7,
marginals very large and sparse.
Appendages.— 60-71. Endites as in L. apus.
Sexual dimorphism. —Normal for the genus.
Reproduction. —Bisexual.
54 A REVIEW OF THE NOTOSTRACA
LEPIDURUS BATESONIsp. nov.
Types. Holotype (<J) and two paratypes in BMNH (1911 .11.8, 23542-4, Norman
collection). Collected by W. Bateson.
Range. Russia. Probably collected at Chilik Kul in the Kazak region, where
Bateson made a collection of other fresh-water entomostraca also in the Norman
collection.
Description. Males only known. Long bodied form in general appearance
(Text-fig. 2 J).
Head. — Eyes and dorsal organ similar to those of Lepidurus lynchi. Second
maxilla present and typical of the genus.
Carapace (Text-fig. 2j). Oval-round, more rounded than in Lepidurus apus.
Carina and its terminal spine absent in all specimens seen ; position of carina demar-
cated only by the light streak of the dorsal blood channel of the carapace which
follows the line of the carina in normal forms. Sulcus wide, rounded, with small sulcal
teeth.
Body. — Segments 33, apodous 8, in all. Apodous ventral marginal spines very
small and widely separated centrally, no supernumeraries.
Appendages. — 49-52. Endites of first thoracic appendage (Text-fig. iod) unique
in known Notostraca ; endites 4 and 5 very short ; much more reduced than in
Lepidurus arcticus and can scarcely have projected beyond carapace margin in
life ; endite 5 of this appendage is little more developed than endite 5 of the second
appendage. Endite 6 of the first appendage is fully developed and claw-like as in
the subsequent appendages (in all other Notostraca the 6th endite of the first thoracic
appendage is reduced to a small soft lobe at the base of the 5th endite).
Telson (Text -fig. 7E-). — Supra- anal plate very similar to that of Lepidurus lynchi,
central spines few (4), marginals few, coarse.
Sexual dimorphism. — Females unknown.
Reproduction. — Bisexual.
Note. — One of the paratypes has an abnormality of the supra-anal plate, which
is reduced in size, soft, and lacks its armature.
references
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A REVIEW OF THE NOTOSTRACA 55
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Mag. nat. Hist. 11 : 496-502.
1924. Some notes on the genus Apus. Ibid. 14 : 559-568.
56 A REVIEW OF THE NOTOSTRACA
Guthorl, P. 1934. Die Arthropoden aus dem Carbon und Perm des Saar-nahe-Pfalz-
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A REVIEW OF THE NOTOSTRACA 57
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30 : 563-567-
ZOOL. 3, T.
PRINTED IN GREAT BRITAIN BY
AD LARD AND SON, LIMITED,
BARTHOLOMEW PRESS, DORKING.
2 SEP 1955
THE POLYCHAETE FAUNA
OF THE GOLD COAST
NORMAN TEBBLE
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 3 No. 2
LONDON: 1955
THE POLYCHAETE FAUNA OF THE
GOLD COAST
BY
NORMAN TEBBLE
Pp. 59-148 ; 1 Chart ; 30 Text-figures
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 3 No. 2
LONDON : 1955
THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, is
issued in five series corresponding to the Departments
of the Museum, and an Historical Series.
Parts appear at irregular intervals as they become
ready. Volumes will contain about three or four
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within one calendar year.
This paper is Vol. 3, No. 2 of the Zoological series.
PRINTED BY ORDER OF THE TRUSTEES OF
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Issued August, 1955 Price One Pound
THE POLYCHAETE FAUNA OF THE
GOLD COAST
By NORMAN TEBBLE
CONTENTS
Page
i. Introduction .......... 61
2. Station Details .......... 61
3. List of Species and Distribution ...... 65
4. Systematic Section ......... 69
1. INTRODUCTION
No large collections of Polychaetes from the Gold Coast have previously been sys-
tematically studied, though a few records were noted by Augener (1918). The
present Faunal Survey results from the examination of littoral collections, down to
a depth of 51 metres, made by Mr. R. Bassindale in 1949-51. Important additional
material has been collected by Mr. J. B. Buchanan. The author is grateful to these
gentlemen for the privilege of studying these collections, through which the known
geographical distribution of many species is extended. Altogether ninety-five
records of Polychaetes from the Gold Coast are here reported, including two new to
science.
In the Systematic Section a key to all families, keys to genera and species where
necessary and a glossary of terms used in the descriptions have been included. All
previous records from the area have been noted although they may refer to species
not examined here. References to species have been restricted, as far as possible,
to standard monographic studies on the group ; the most important are Augener,
1918, and Fauvel, 1923 and 1927a.
Dr. A. B. Hastings kindly identified the Polyzoan specimens with which some
Serpulidae are associated and the author also wishes to acknowledge the technical
assistance he has received from Miss A. C. Edwards, of the British Museum (Nat.
Hist.) staff, particularly in the drawing of figures and charts.
2. STATION DETAILS
Littoral collections were made at the following places ; the details have been
provided by Mr. R. Bassindale.
I. Accra (5 30' N., o° 15' W.) (a) Christiansborg. The reef at Christiansborg
lies at the foot of the Governor's Castle (2 miles east of Accra lighthouse)
zool. 3, 2. 6
62
THE POLYCHAETE FAUNA OF THE GOLD COAST
and consists of native rock with small deep pools mainly at the lowest
levels. There is sand and just a little exposed rock above mid-tide level.
(b) Chorkor. A seine net was operated from the sand beach here, two
miles west of Accra lighthouse.
II. Tenpobo and Pram Pram, 24 and 23 miles, respectively, east of Accra. The
coast is of sand and the upper three-quarters of the beach is a steep sand
slope. Near the headlands of a series of shallow bays there are low level
rocks and boulder reefs jutting out to sea. These reefs occupy the lowest
foot of spring tide range except for native rock, which may rise one or
two feet above this level. Maximum tidal range is six feet, and neap
tides do not expose the levels from 0-1 foot above chart datum (low tide
level of ordinary spring tides) . At Tenpobo the reefs are protected from
low level wave wash by a line of high rocks to seaward. There are many
small boulders and stones on the reef ; gravel and sand packing between
the stones is common.
III. Winneba, 32 miles west of Accra. The beach here is of native rock from below
low tide level to above high tide level.
IV. Apam, 39 miles west of Accra. The shore is of rock and boulders set in sand
with rock pools. Extending to the east is a shallow sand bay with sandy
mud in the shelter of the Apam headland.
V. Sekondi, 113 miles west of Accra. Collections here were made from the hori-
zontal surface of a rock plateau at or above high tide level. The waves
wash over the plateau at most states of the tide and so replenish the pools.
There is probably no period of complete submergence of the plateau.
VI. Dixcove, 131 miles west of Accra. The bay at Dixcove is almost square, the
sides being about 400 yards long. The innermost side of the bay has a
sandy beach with large boulders and native rock at low tide level —
the rocks rising to mid-tide level or higher. Collections were also made
on the western side of the bay which consists of small boulders and
stones, with sand, mud and gravel closer in.
VII. Princes Town, 142 miles west of Accra. A solid rocky headland very much
wave-washed, with a few boulders round the headland to the east.
VIII. Axiwi, 154 miles west of Accra : (a) The Hospital Reef consists of a narrow
belt of rock, boulders and stones jutting out to sea. Collections here
were made from the exposed side.
(b) The Lighthouse Reef juts out in a similar way but is protected by
two islands, one half way along and one at the seaward end. In-shore
the reef is at low level and has stones with gravel, sand and mud. Further
out the stones increase in size until very large boulders occur on the
exposed side. Collections here were made mainly from the sheltered
side.
Off-shore collections were made by Mr. Bassindale entirely within the area off
Accra shown on the chart (p. 63). The following lists give further details of the
stations. Several collections were made by Mr. Bassindale at stations which are not
THE POLYCHAETE FAUNA OF THE GOLD COAST
6.3
6 4
THE POLYCHAETE FAUNA OF THE GOLD COAST
numbered and they are referred to in the text by reference to points on the coast.
Mr. Buchanan made part of his survey within the area indicated on the chart between
3*6 and n-o metres ; other stations at which he made collections have also been
referred to as the Buchanan Survey and their positions have been indicated in the
same way.
List of Collecting Stations in the Sea off Accra
All dredge hauls of 10 minute duration and Agassiz Trawl hauls of 15 minute
duration except where stated.
Station
Depth in
number
Date
metres
Instrument used
5
9.xi.5o
13
Large dredge
6
,,
12
» >»
10
19.xi.50
14
9 min
11
,,
13
>
12
,,
16
»
14
28.xi.50
20
»
15
,,
16
»
16
,,
17
»
27
14.xii.50
22
5 min
28
,,
30
8 min
29
20.xii.50
13
>
32
,,
11
,
33
,,
28
»
35
21.xii.50
37
30 min
44
31.xii.50
35
,
47
4-i-5i
44
Small dredge 15 min
48
44
Large dredge
52
45
Small dredge
53
i5-i-5i
11
.
54
11
> >>
55
12
• >>
56
16
• > >
57
i7-i-5i
16
,,
58
20
•
59
24
,,
61
18.L51
26
,,
62
30
>>
65
22.L51
7
>,
66
10
, ,
67
x 4
,,
68
19
,,
69
,,
22
,,
70
24.i.5i
25
>>
7i
30
,,
73
4i
,,
80
21.ii.51
10
Agassiz Trawl
88
5-iii-5i
13
,,
89
7-iii-5i
16
THE POLYCHAETE FAUNA OF THE GOLD COAST
65
List of Collecting Stations in the Sea off Accra — cont.
Station
Depth in
number
Date
metres
Instrument used
92
12.iii.51
n
Agassiz Trawl
93
>>
12
94
>»
17
95
„
17
„ 30 mm
97
14.u1.51
20
99
>»
28
104
29.u1.51
13
no
4.IV.51
40
20 mm
in
,,
43
115
5iv.5i
24
No details
121
11.iv.51
8
Agassiz Trawl
123
>
9
124
12. iv. 51
11
125
,,
16
126
,,
20
127
14. iv. 51
17
130
26.iv.51
32
131
2.V.51
37
132
>*
44
133
»>
5i
Approximate distance covered in normal 10-minute haul with large dredge was 700 metres,
in normal 15 -minute haul with Agassiz Trawl, 400 metres. (More in shallow and less in deep
water.)
3. LIST OF SPECIES AND DISTRIBUTION
X = Collected in the present survey.
o = Recorded by Augener (1918), not collected in the present survey.
Localities
A
Species
Aphroditidae
Aphrodite alta
Hermione hystrix
Panthalis bicolor
Euthalenessa insignis
Eusigalion vazensis
Sthenelais boa .
limicola
Leanira japonica
,, yhleni
a5
o3
3
a
<u
pj
o3
t— 1
O
$
Ph
<
<
tf
CO
,__,
3
rt
+->
A
Ph
rS
O
tuO
X
^
66
THE POLYCHAETE FAUNA OF THE GOLD COAST
List of Species and Distribution — cont.
X = Collected in the present survey.
o = Recorded by Augener (1918), not collected in the present survey.
Localities
Species
Scalisetosus pellucidus
Acholoe astericola
Pareulepis geayi
Lepidonotus hupferi .
Harmothoe goreensis
5 g
O o3
H
rf
<u
aj
.'2
O
.£5
<u
c
O
&
s
o
Chrysopetalidae
Bhawania goodei
Amphinomidae
Chloeia viridis .
Hermodice carunculata var didymo-
branchiata .
Eurythoe complanata
parvecarunculata
Phyllodocidae
Phyllodoce africana .
oculata .
Eteone picta
Hesionidae
Leocrates claparedii .
Syllidae
Trypanosyllis prampramensis
Syllis gracilis .
variegata
„ hyalina .
,, cornuta .
Opisthosyllis brunnea
Pionosyllis ehlersiaeformis
Exogene gemmifera .
Nereidae
Nereis falsa
Neanthes succinea
Perinereis melanocephala .
Pseudonereis gallapagensis
X
XX X
. . X
x . . X
. . . . X
THE POLYCHAETE FAUNA OF THE GOLD COAST
67
List of Species and Distribution — cont.
X = Collected in the present survey.
o = Recorded by Augener (19 18), not collected in the present survey.
Localities
Species
Nephthydidae
Nephthys hombergii .
Aglaophamus lyrochaetus
Glyceridae
Glycera convoluta
Goniada multidentata
Goniadopsis incerta .
Ophioglycera archeri sp. n.
EUNICIDAE
Eunice antennata
,, vittata .
filamentosa
coccinea
gracilis
rubra .
Marphysa dartevellei
Lysidice ninetta
collaris
Diopatra musseraensis
,, neapolitana
Onuphis eremita
Lumbrineris impatiens
,, albifrons
Holla parthenopeia .
Orbiniidae
Orbinia foetida var linguistica
Scoloplos madagascarensis
,, dubia, sp. n.
Spionidae
Prionospio pinnata .
ClRRATULIDAE
Audouinia tentaculata
,, filigera .
punctata .
Cirratulus filiformis .
T3
a
d
o
o
0)
H
8
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68
THE POLYCHAETE FAUNA OF THE GOLD COAST
List of Species and Distribution — cont.
X = Collected in the present survey
o = Recorded by Augener (1918), not collected in the present survey.
Localities
Species
Flabelligeridae
StylaHoides scutigeroides .
„ arenosus
„ tropicus
Opheliidae
Armandia intermedia
Polyophthalmus pictus
Maldanidae
Maldane sarsi
OWENIIDAE
Owenia fusiformis .
Sabellariidae
Sabellaria eupomatoides
spinulosa var intoshi
,, spinulosa var alcocki
Sternaspidae
Sternaspis scutata var africana
Ampharetidae
Phyllamphicteis collaribranchis
Pterolysippe bipennata
Isolda whydahensis .
Terebellidae
Pista grubei
Terebellides stroemi
Streblosoma persica
Loimia medusa
Amaea accraensis
Sabellidae
Laonome puncturata
Dasychone lucullana
Potamilla casamancensis
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THE POLYCHAETE FAUNA OF THE GOLD COAST
69
List of Species and Distribution— cont.
X = Collected in the present survey
o = Recorded by Augener (1918), not collected in the present survey.
Localities
„ , — .> . — .
Species
Serpulidae
Salmacina incrustans
Hydroides norvegica
spinosus .
,, arnoldi
Apomatus similis
Serpula vermicularis
Vermiliopsis multicristata
prampramiana
Spirorbis sp. .
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Augener (191 8) from the Cape Coast Castle.
4. SYSTEMATIC PART
(a) Key to Families of Polychaeta
The following key has been adapted from Fauvel (1923 and 1927a). It includes
families not collected in the present survey and these have been marked with an
asterisk.
1. Some parapodia bearing elytra, the rest with dorsal cirri . . . Aphroditidae.
Parapodia without elytra . . . . . . . . . . 2.
2. A fan-shaped group of broad, flattened chaetae (paleae) on all segments
Chrysopetalidae.
No such groups of chaetae .......... 3.
3. Prostomium not distinct ; when present pedal cirri globular . . . . 4.
Prostomium distinct ......... .6.
4. Parapodia biramous and achaetous ; prostomium fused with the two succeeding
segments and bearing 2 or 4 tentacular cirri containing aciculae ; pedalcirri absent.
Pelagic ........... Tomopteridae.*
Parapodia uniramous, with chaetae and with globular cirri ..... 5.
5. Pharynx armed .......... Pisionidae.*
Pharynx unarmed ......... Sphaerodoridae.*
6. Body not normally divided into distinct regions ; all segments alike except those
near the mouth and the pygidium. Pharynx protrusible and normally armed
with jaws. This group includes those families of the Polychaeta Errantia of
Fauvel (1923) not given above ......... 7.
7 o THE POLYCHAETE FAUNA OF THE GOLD COAST
Body often divided into distinct regions and adapted to either a tubicolous or
burrowing life, the head is, therefore, either greatly modified, with a wreath of
tentacles or gills surrounding the mouth, or almost completely divested of cephalic
appendages. Parapodia much modified in the tubicolous forms in which the
neuropodia are often in the form of pinnules (tori) bearing hooks or uncini.
Without jaws. This group includes the Polychaeta Sedentaria of Fauvel,
1927a 15-
7. Prostomium conical, without antennae or palps ; dorsal and ventral cirri foliaceous ;
chaetae rare, when present simple, acicular. Pelagic . . Typhloscolecidae.*
Prostomium with tentacles, usually with palps . . . . . . . 8.
8. Prostomium small with three antennae ; pharynx unarmed ; a caruncle almost
always present. Chaetae simple, calcareous in some genera . Amphinomidae.
Prostomium prominent ; without a caruncle . . . . . . . 9.
9. Prostomium long, conical, annulated, with four short terminal antennae Glyceridae.
Prostomium not so ........... 10.
10. Proboscis without chitinous teeth . . . . . . . . .11.
Proboscis with chitinous teeth . . . . . . . . . .14.
11. Parapodia normally uniramous, cirri usually foliaceous . . . . .12.
Parapodia biramous or very much reduced . . . . . . 13.
12. Prostomium with two very large globular eyes. Pelagic . . . Alciopidae.*
Prostomium with two small eyes ....... Phyllodocidae.
13. Parapodia biramous, with inter-ramal cirri (branchiae) . . . Nephthydidae.
Parapodia biramous, or very much reduced, without branchiae . . Hesionidae.
14. Proboscis with one pair of heavy curved jaws distally and normally with paragnaths
on its dorsal and ventral surfaces ........ Nereidae.
Proboscis with a complex armature — a pair of ventral pieces and two, or frequently
more, dorsal pieces ......... Eunicidae.
Proboscis with one large tooth, or crown of teeth, or both .... Syllidae.
15. Body clearly divided into distinct regions . . . . . . . 23.
Body not clearly divided into distinct regions . . . . . . 16.
16. Body very much swollen, few segments, filiform anal branchiae. A large ventral
shield bordered with chaetae ....... Sternaspidae.
Body not swollen, numerous segments, without anal branchiae. No ventral
shield ............. ij.
17. Palps elongate ............ 18.
Without palps ............ 22.
18. Elongate palps (tentacular cirri) on the body, not on the prostomium . Cirratulidae.
Elongate palps on the prostomium . . . . . . . . ig.
19. A cephalic cage formed by the chaetae of the first chaetigers . Flabelligeridae.
Without a cephalic cage . . . . . . . . . . .20.
20. Without branchiae. Prostomium oval, spoon shaped. Palps with sucker-like
papillae ........... Magelonidae.*
With branchiae ............ 21.
21. Parapodial lamellae erect, dorsal. Branchiae cirriform. Hooded hooks and
capillary chaetae .......... Spionidae.
Anterior parapodial lamellae frilled or flask-shaped. Branchiae filliform. Chaetae
of many kinds, plumed, spiny, barbed and acicular, etc. . . . Disomidae.*
22. One median antenna. Dorsal foliaceous branchiae. Hooded hooks and capillary
chaetae ........... Paraonidae.*
Two antennae or none. Capillary and forked chaetae, no hooded hooks
SCALIBREGMIDAE.*
Prostomium blunt, without appendages or with a crown of lobes. No branchiae.
Ventral tori with rows of uncini. Sandy tube ..... Oweniidae.
Prostomium without appendages, with a keel or rimmed cephalic plate . Maldanidae.
THE POLYCHAETE FAUNA OF THE GOLD COAST
V-
23-
24.
25-
26.
27
28.
29.
30-
31
32.
A terminal branchial tuft with numerous filaments. Thoracic uncini dorsal
abdominal uncini ventral ........
No terminal branchial tuft ........
Paleae forming an operculum to close the tube .....
No opercular paleae .........
Prostomium without processes. Branchiae on many segments, but not on the
prostomium . . . . . .
Prostomium with processes, either branchiae or tentacular filaments
With uncini ...........
Without uncini ..........
With acicular hooks or capillary chaetae ......
With no acicular hooks, only capillary chaetae .
Prostomium blunt. Non-retractile arborescent
branchiae in the
32
24
31
25
26
29
28
27
Orbiniidae
Opheliidae
middle body
region ........... Arenicolidae.*
Prostomium conical. Branchiae, if present, posterior in position, simple or multifid,
if multifid retractile ......... Capitellidae.*
Body divided into 2-3 strikingly dissimilar regions. Prostomium with or without
2 small tentacles. Two long grooved palps. Anterior region short with uni-
ramous feet bearing peculiar chaetae in the 4th chaetiger. Posterior notopodia
erect. Uncini comb-like ....... Chaetopteridae.*
Body divided into regions which are not strikingly dissimilar. With numerous
tentacular filaments . . . . . . . . . . 30.
Ampharetidae.
Terebellidae.
Amphictenidae.
Sabellariidae.
Serpulidae.
Sabellidae.
Tentacular cirri retractile into the mouth. Prostomium distinct .
Tentacular cirri not retractile into the mouth. Prostomium indistinct
An operculum of one anterior row of golden paleae
An operculum of two stalks bearing a crown of paleae
Tube calcareous, normally with an operculum ....
Tube membranaceous or mucous, never with an operculum .
(b) Glossary of Terms
The following glossary includes only those terms which are not normally to be
found in a standard text-book of zoology or English Dictionary.
Acicula : A stout chaeta of one piece, almost always tapering to a blunt point.
Capillary Chaeta : Simple chaeta, normally long and slender.
Caruncle : A longitudinal ridge on the dorsal surface of the prostomium and sometimes
extending to some of the following segments.
Chaeta : A bristle made largely of chitin but exceptionally some may be composed of calcium
carbonate (e.g. Eurythoe complanata). Chaetae are either simple (of one piece), or com-
pound (having two or more articulating pieces).
Falciger : The terminal article of a compound chaeta when it is short and broad. c.f.
Spiniger.
Heterogomph : The condition when the two lateral branches of the articulating surface of
the main stem of a compound chaeta are unequal in length.
Homogomph : The condition when the two lateral branches of the articulating surface of the
main stem of a compound chaeta are equal in length.
Limbate : The condition when a simple chaeta is bordered along all or part of its length, e.g.
Apomatus similis, Text-fig. 30, /.
Palea : A stout simple chaeta, groups of which appear in the Chrysopetalidac, Amphictenidae,
Ampharetidae and Sabellariidae.
Sesquiramous : Condition of a parapodium when the dorsal component (notopodium) is
reduced to a few chaetae or an acicula.
72 THE POLYCHAETE FAUNA OF THE GOLD COAST
Spiniger : The terminal article of a compound chaeta when it is long and slender, c.f.
Falciger.
Stylode : A dorsal appendage of the branchial filaments in some Sabellidae.
Uncinus : A short simple chaeta which may be sigmoid, with teeth at the tip, or plate-like
with rows of teeth along one edge.
(c) Descriptions of Species
Keys to genera and species have been compiled only in respect of forms actually
recorded from the Gold Coast, or where systematic considerations suggest that details
of related forms will be of help to the student.
Examples of all species have been deposited in the collections of the B.M. (N.H.),
Reg. Nos., 1953.3.1.1-1234 ; duplicate material covering the majority of species,
has been deposited in the Zoology Dept., University College, Achimota, Gold Coast.
Family Aphroditidae
Prostomium normally with four sessile, or pedunculate, eyes, one to three antennae
and two palps inserted ventrally. Protrusible proboscis crowned with papillae
and with four chitinous jaws (except the Aphroditinae) . A feature of this family
is the possession of elytra on the dorsal surface of the parapodia. The first pairs
are always attached to segments 2, 4, 5 and 7, thereafter the arrangement varies,
posteriorly they may be absent. Parapodia biramous, exceptionally reduced.
Key to Subfamilies
1. Body short, fat and oval, jaws absent or rudimentary. Elytra may be covered with
a thick mat of felt ......... Aphroditinae.
Body comparatively long and thin, pharynx with jaws. Elytra never covered with
a mat of felt ............. 2.
2. Compound chaetae in the neuropodia. Elytra on all segments posteriorly Sigalioninae.
All chaetae simple. Elytra on every second or third segment, or entirely absent,
from the posterior part of the body . . . . . . . . . 3.
3. Elytra on every second segment in the posterior part of the body. Prostomial eyes
sessile or stalked. Tubicolous in habit ...... Polyodontinae.
Elytra on every third segment in the posterior part of the body, or entirely absent.
Prostomial eyes sessile, never stalked. Errantiate in habit . . Polynoinae.
Subfamily Aphroditinae
One unpaired antenna, no laterals ; a facial tubercle beneath the antenna.
Elytra 15 pairs, rarely 20, inserted on segments 2, 4, 5, 7, 9-21, 23, 25, 28, 31,
alternating with dorsal cirri.
Key to Genera
1 . Dorsal surface covered with a thick felt ; dorsal notopodial chaetae ending in blunt
points or with fine hooks ......... Aphrodite.
2. No covering of felt ; notopodial chaetae harpoon-shaped on elytrigerous segments
Hermione.
THE POLYCHAETE FAUNA OF THE GOLD COAST 73
Genus APHRODITE Linnaeus, 1758
Fifteen pairs of elytra. Eyes sessile or absent.
Aphrodite alt a Kinberg, 1856
Aphrodite alta, Mcintosh, 1924, pp. 5-6, from Cape of Good Hope waters.
Aphrodite alta, Monro, 1930, pp. 36-38, fig. $a-i, from 64 20' 00" S., 63 01 ' 00" W.
Localities. Off Accra : Stn. 47 (1) ; Stn. no (1) ; Stn. 133 (1) ; all complete,
20 mm. long by 10 mm. wide.
This species is characterized by the lack of eyes, the dorsal notopodial chaetae
which taper to a fine hooked point, and which appear to lie in the dorsal felt rather
than to project through it, the thick bearded ends of the neuropodial chaetae and
the posterior neuropodial chaetae with long alternating teeth. These features
clearly distinguish the species from A. aculeata which Fauvel (1950) reported from
Rio de Oro.
Distribution. This appears to be the first record of A. alta from the W. coast
of Africa, north of the Cape. Kinberg (1856) reported it from Rio de Janeiro.
Genus HERMIONE Blainville, 1828
Four pedunculate eyes. Fifteen pairs of elytra.
Hermione hystrix (Savigny), 1818
Hermione hystrix, Fauvel, 1923, pp. 35-36, fig. 11.
Locality. Off Accra, Stn. 126 (12), all complete, measuring between 20-
30 mm. long by 10-15 mm. wide.
There are two pairs of eyes, each pair carried on a peduncle, situated ventral to
the antenna. Two types of notopodial chaetae are present on elytrigerous segments ;
one set, stout, curved and ending in a simple point, the others with strong, harpoon-
shaped points. On cirrigerous segments there are only thin chaetae with simple
points. Neuropodial chaetae are bifurcate with a short limb and a long curved one
which may be toothed on anterior and posterior feet.
Distribution. Mediterranean Sea, Atlantic Ocean and Indo-Pacific regions.
Subfamily Polyodontinae
Bilobed prostomium, with four sessile eyes, or two large ommatophores and three
antennae. Elytra on segments, 2, 4, 5, 7, 9, etc.
Only one genus of this subfamily has been recorded from the Gold Coast, Pan-
thalis, but the following Key is included here to illustrate the difference between
it and Polyodontes.
1 . Superior neuropodial chaetae elongate hastate, the tip smooth or more or less hirsute
Polyodontes.
2. Superior neuropodial chaetae penicillate (Text-fig. 1) . . . . Panthalis.
74 THE POLYCHAETE FAUNA OF THE GOLD COAST
Hartman (1939) has drawn attention to the need for revising the diagnostic generic
characters in this subfamily and this Key has been adopted from her suggestions.
Thus the presence or absence of branchial lobes is of doubtful value, for many speci-
mens are incomplete when collected and the methods adopted for preservation fre-
quently result in the complete retraction of these organs.
Genus PANTHALIS Kinberg, 1855
Panthalis bicolor Grube, 1877
Polyodontes bicolor, Augener, 1918, pp. 119-125, pi. 2, fig. 28 ; pi. 3, fig. 47 ; text-fig. 4. From
Accra.
Localities. Off Accra, Stn. 33 (1), anterior fragment 23 mm. in length ; Buchanan
Survey, in a depth of 3-6 to n -o metres (1), anterior fragment, 10 mm. long ; Buch-
anan Survey, in a depth of 10 metres off lighthouse (1), anterior fragment, 17 mm.
long.
The prostomium carries two large anterior ommatophores (eye-stalks) and two
small posterior eye-spots, situated on either side of the median antenna, which
projects forward between the ommatophores. Two lateral antennae are ventral
to the eye-stalks. There are two pairs of tentacular cirri. Spinning glands are
present in the feet. Neuropodial chaetae are illustrated in Text-fig. 1, (a) superior
penicillate with the tip frequently carried beyond the bushy portion, (b) median
stout aristate, (c) inferior serrulate.
No tubes were found with these specimens.
Distribution. The species is known only from the tropical West African coast.
Subfamily Sigalioninae
Elytra numerous, attached to segments 2, 4, 5, 7, etc., and on each segment from
the 23rd-29th up to the end of the body. All the genera here reported have three
antennae ; other genera, however, have one (Pholoe) or two (Sigalion).
Key to Genera
1. With a dorsal cirrus on the 3rd chaetiger . . . . . . . . 2.
Without a dorsal cirrus on the 3rd chaetiger ....... 3.
2. With the unpaired antenna inserted on the anterior margin of the prostomium
Enthalenessa.
With the unpaired antenna inserted on the middle of the prostomium, in line with,
or posterior to, the eyes .......... Eusigalion.
3. With falcigerous compound chaetae in the neuropodium .... Sthenelais.
With spinigerous compound chaetae in the neuropodium .... Leanira.
Genus EUTHALENESSA Darboux, 1899
The median unpaired antenna, without ceratophore or ctenidia, is on the anterior
margin of the prostomium. From the fourth chaetiger dorsal ctenidia are present
on the parapodia ; on the third foot there is a branchial tubercle but no true dorsal
cirrus.
THE POLYCHAETE FAUNA OF THE GOLD COAST
75
Fig. i. Panthalis bicolov. Neuropodial chaetae ; (a) superior penicillate ; (b) median
aristate ; (c) inferior serrulate, x 200. (Specimen from the Buchanan Survey.)
Euthalenessa insignis Ehlers, 1908
Euthalenessa insignis Augener, 1918, pp. 108-112, from Apam.
This species has not been found in the present collections.
The specimens described by Augener were characterized by the possession of a
third cirrus on the first chaetiger, and it is doubtful if this character is applicable
to Ehler's species, but specimens will have to be examined before the true identity
of Augener's record can be established.
Genus EUSIGALION, Augener, 1918
The median antenna is in the middle of the prostomium, between, or posterior to,
the eyes. This character is here taken to separate Eusigalion from Euthalanessa,
which are similar in other respects.
ZOOL. 3, 2. 7
7 o
THE POLYCHAETE FAUNA OF THE GOLD COAST
Eusigalion vazensis Augener, 1918
Eusigalion vazensis Augener, 1918, pp. 113-118, pi. 2, fig. 14 ; pi. 3, figs. 44-46 ; text-fig. 3.
From Fernand Vaz in the French Congo.
Locality. Off Accra, Buchanan survey, in 3-6 to n-o metres depth (2), anterior
pieces, 20-23 mm. long.
There are about 12 pinnate filaments on the outside edge of each elytron. An-
tennae are very short, the unpaired median is set back on the dorsal surface of the
prostomium, between the two pairs of small eyes ; the paired anterior antennae
project from the front border of the prostomium. Notopodial simple chaetae are
long, barred and serrated. Neuropodial chaetae are in two groups ; a supra-
acicula group of simple spinose bristles and long-shafted compound forms with
numerous joints, and a sub-acicula group, all compound, one set with single long-
shafted, bifid, terminal articles, the other with numerous articles of which the
terminal one is also bifid.
Distribution. Only known from the above records.
Genus STHENELAIS Kinberg, 1855
Prostomium with four eyes. The lateral antennae are fused with the first chaetiger.
Unpaired median antenna with basal ctenidia. Two long palps with ctenidia at the
base. Branchiae on all parapodia from the fourth. Notopodial chaetae simple ;
neuropodial chaetae compound f alcigers with sometimes a superior group of simple
bristles.
Key to Species
1 . Elytra with simple fringes and entire borders ......
Anterior elytra with bifid fringes ;
border notched
S. boa.
elytra from the 24th chaetiger with outside
5. limicola.
Sthenelais boa (Johnston), 1833
Sthenelais boa, Fauvel, 1923, pp. 110-111, fig. 41, a-l.
Locality. Tenpobo shore, 6.U.50 (1), 43 mm. long.
Elytra normally reniform, crossing and overlapping on the back, with minute
papillae and with simple fringes on the outer border. Notopodial chaetae simple,
finely spinous. Superior neuropodial chaetae simple, bi-pectinate, strongly spinous,
the middle group compound f alcigers with single bidentate articles, the inferior group
also compound with long, pseudo-multiarticulate, bidentate falcigers.
Distribution. English Channel, Mediterranean Sea, Atlantic Ocean, Indian Ocean.
Sthenelais limicola (Ehlers), 1864
Sthenelais limicola, Fauvel, 1923, pp. 113-114, fig. 42, a-g.
Locality. Off Accra, Buchanan Survey, in 3-6-11-0 metres depth (4), two com-
plete, 60, 75 mm. in length, two fragments 23 and 56 mm. long.
THE POLYCHAETE FAUNA OF THE GOLD COAST 77
The characters noted in the above key serve to separate this species from S. boa ;
in other characters the species are similar.
Distribution. North Sea, Mediterranean Sea, Atlantic Ocean.
Genus LEANIRA Kinberg, 1855
Members of this genus may, or may not, have eyes ; otherwise they are similar
to Sthenelais, except as keyed above.
Key to Species
1 . Superior neuropodial chaetae simple, with whorls of spikes ; inferior neuropodial
chaetae compound, caniculate spinigers . . . . . . L. japonica.
2. All neuropodial chaetae compound, caniculate spinigers . . . L. yhleni.
Leanira japonica Mcintosh, 1885
Sthenolepis japonica, Willey, 1905, pp. 259-260, pi. 2, rig. 49.
Leanira japonica, Fauvel, 1932, pp. 33-34.
Localities. Off Accra, Stn. 57 (1) ; Stn. 70 (3) ; Stn. 97 (1) ; Stn. 132 (1) ; Stn.
I 33 ( 2 )- One of the specimens from Stn. 70 is complete, 64 mm. in length. All
others are fragmentary, up to 15 mm. long.
Prostomium with four eyes. The dorsal cirrus is replaced in this species by a
very small tubercle. The elytra, which are fringed, overlap but do not cross, and
leave a considerable area of the mid-dorsal surface exposed. Notopodial chaetae
are long and slender, neuropodial compound chaetae have long caniculate terminal
pieces and a superior group of simple chaetae, with whorls of spikes, on most
parapodia.
Distribution. Previously reported from Japan, Malaya, Bay of Bengal and the
Arabian Sea, this appears to be the first record of this species from the Atlantic
Ocean.
Leanira yhleni Malmgren, 1867
Leanira yhleni, Fauvel, 1923, pp. 117-118 (no figures).
Locality. Off Accra, Buchanan Survey, 41*4 metres depth, off Lagoon, (1),
anterior fragment 25 mm. long.
This species is distinguishable from L. japonica as noted above in the key, and in
having the anterior pair of eyes larger than the posterior pair. In L. japonica the
posterior eyes are larger than the anterior. In both species the anterior eyes are
near the front border of the prostomium, partly hidden beneath the antennal ctenidia,
and the posterior pair are at the base of the unpaired antenna.
Distribution. Bay of Biscay, Mediterranean Sea, Adriatic Sea ; this appears
to be the first record from the equatorial Atlantic.
Subfamily Polynoinae
Elytra 12 or more pairs inserted on segments 2, 4, 5, 7, 9, to 23-26, thence on every
third segment to the posterior end of the body. All genera here reported have one
unpaired, median antenna and two paired, lateral antennae.
78 THE POLYCHAETE FAUNA OF THE GOLD COAST
Key to Genera
i. Lateral antennae inserted ventrally ; 15 pairs of elytra . . . . . 2.
Lateral antennae inserted terminally, continuous with the prostomial peaks . . 3.
2. Chaetae transparent as crystal, with spinous pouches .... Scalisetosus.
Chaetae not transparent, without spinous pouches ..... Harmothoe.
3. Elytra numerous, more than 15 pairs, a large dorsal tubercle in the form of a T on
the cirrigerous segments ......... Acholoe.
Elytra 12 pairs ............ 4.
4. Segment 3 fused with the segments 2 and 4 dorsally ..... Pareulepis.
Segment 3 not fused dorsally ........ Lepidonotus.
Genus SCALISETOSUS Mcintosh, 1885
The 15 pairs of elytra are inserted on segments 2, 4, 5, 7-23, 26-29-32, not covering
the whole body. Neuropodial chaetae with semilunar cusps are slightly bifid at the
extremity. Notopodial chaetae curved with blunt spines.
Scalisetosus pellucidus (Ehlers), 1864
Scalisetosus pellucidus, Fauvel, 1923, p. 74, fig. 27, a-f.
Locality. Off Accra, stn. 47 (1), 10 mm. in length.
Irregular dark brown transverse bands decorate the dorsal surface. Antennae
and cirri are papillated. The ventral chaetae are not as deeply incised at their distal
ends and the notopodial chaetae not so clearly notched as normal representatives
of the species.
Distribution. Mediterranean Sea, Atlantic Ocean, Bay of Bengal, Malay
Archipelago.
Genus ACHOLOE Claparede, 1870
Two pairs of tentacular cirri with basal aciculae. Elytra on segments 2, 4, 5, 7-23,
26, 29, 32, thence almost up to the end of the body. Parapodia with reduced noto-
podia.
Acholoe astericola (Delle Chiaje), 1823
Acholoe astericola, Fauvel, 1923, pp. 94-95, fig. 36, d-h.
Localities. Off Accra : From starfish, at 25 metres, and 10 metres (off Petrol
Buoy), numerous specimens; Stn. 57 (1), among tubes of Diopatra neapolitana ;
Stn. 67 (1) ; Stn. 80 (1) ; Stn. 89 (1) ; Stn. 104 (1) ; Stn. 123 (1).
Most of these specimens are fragmentary, the complete ones measure up to 30 mm.
long.
There are about 45 pairs of elytra. Neuropodial chaetae have a recurved un-
identate tip with a short spinous region, and notopodial chaetae are short with trans-
verse rows of pinnules.
Distribution. English Channel, Mediterranean Sea, Atlantic Ocean.
THE POLYCHAETE FAUNA OF THE GOLD COAST
79
Genus PAREULEPIS Darboux, 1899
Segment 3 fused with segments 2 and 4 dorsally. Elytra on segments 2, 4, 5, 7,
>, 11-21-24. Notopodial chaetae are slender capillaries and stout curved bristles
>ent abruptly at their distal ends. Neuropodial aciculae are short, wholly embedded,
vith extended, plate-like terminations. Neuropodial chaetae include a few small,
aterally pectinate, bristles, and a bundle of almost straight chaetae.
Pareulepis geayi (Fauvel), 1918
lulepis geayi, Fauvel, 1919, PP- 335-339, pi- 15, figs. 17-21 ; pi. 17, figs. 76-79.
Locality. Off Accra, Buchanan Survey, between 11-13 metres depth, off
'hristiansborg Castle (2), each about 25 mm. long.
The arrangement of anterior segments in this species is shown in Text-fig. 2. Thus
egment 3 is represented laterally by parapodia, but is fused dorsally with segments
SEGMENT
2
SEGMENT
3
SEGMENT
4
Fig. 2. Pareulepis geayi. Dorsal view of anterior segments to show the fusion of
segment 3 with segments 2 and 4 dorsally. Elytra and the 2nd and 3rd parapodia on
the left-hand side have been omitted.
5 and 4. Elytra are present therefore on segments 2, 4, 5, 7, 9-21-24, after which
here are ten cirrigerous segments up to the end of the body. The elytra on segment
*4 extends from segment 23 backwards to 29, so that only a small posterior portion
>f the worm remains uncovered. There are up to 12 marginal papillae on the outside
border of each elytron. Dorsal cirri are stout and foliaceous throughout.
The specimens here described have been referred to this species, and not to P.
Imbriata Treadwell, with which Hartman (1939) considers P. geayi synonymous,
80 THE POLYCHAETE FAUNA OF THE GOLD COAST
because of the position of the unpaired median antenna. Fauvel (1919, p. 338)
clearly implies that the unpaired antenna is posterior to the median line of the pro-
stomium. Treadwell (1900, p. 190), however, places it on the anterior prostomial
margin. Too, Treadwell's specimens had eye-spots ; these are not present in Fauvel's
species.
Distribution. This species appears to have been recorded only from the Indo-
Pacific region, though the genus, through P.fimbriata, is also known from the Pacific
coast of America.
Genus LEPIDONOTUS Leach, 1816
Prostomium bilobed, without frontal horns, four eyes. Twelve pairs of elytra
attached to segments 2, 4, 5, 7-21, 23.
Lepidonotus hupferi Augener, 1918
Lepidonotus hupferi Augener, 1918, pp. 133-136, pi. 2, figs. 7-1 1 ; text-fig. 5.
Localities, (i) Tenpobo shore (10), most complete, between 8-18 mm. in length,
collected in February, 1950. (2) Off Accra, from wood netted off Chorkor in 14
metres, 14.V.51 (1), 10 mm. long. (3) Axim, (a) Shore, 13/14.^.49 (1), 12 mm. long ;
{b) Hospital Reef (4), 2 complete, 4 and 6 mm. long ; 2 parts 4 and 8 mm. long.
The elytra completely cover the dorsal surface, are fringed on their external
margins, and covered with very small tubercles, with a broad base and slender stalk.
Notopodial chaetae are slender and serrated. Neuropodial chaetae are stout with a
serrulated blade ; some are distally entire, others have a slightly bifid tip.
Fauvel (1950) has noted that this species appears to replace L. squamatus in the
tropical zone.
Distribution. Tropical West Africa, Western Mexico, Panama, N.W. South
America. Hartman (1939) records this Western Hemisphere distribution from the
intertidal zone to 22 metres.
Genus HARMOTHOE Kinberg, 1855
Prostomium bilobed, with four eyes, often with lateral peaks. Elytra covering
all the dorsal surface on segments 2, 4, 5, 7-23, 26, 29, 32.
Harmotho'e goreensis Augener, 1918
Harmothoe goreensis Augener, 1918, pp. 142-146, pi. 2, figs. 4-6 ; pi. 3, fig. 42 ; text-fig. 7.
Localities. Off Accra : (a) Stn. 47, (1), 8 mm. long ; (b) from wood netted off
Chorkor in 14 m., (4) ; (c) 2 miles out from R. Densu, 8.iv.49, 14-6 m., in shingle,
(4), 2 complete, 6 mm. long ; (d) 2 miles west out from R. Densu, 1 mile off shore,
2.iii.49, 7-3 m., (1), 4 mm. long.
The elytra have short fringes on their external margins ; elytron tubercles are
sac-like in shape with a minute distal pore. Tentacular cirri, dorsal cirri and palps
are covered with papillae. Neuropodial chaetae are normally bidentate superiorly
THE POLYCHAETE FAUNA OF THE GOLD COAST 81
and inferiorly are unindentate. Notopodial chaetae (always stouter than neuro-
podial bristles in Harmothoe) are distally entire and heavily serrated.
Distribution. Previously recorded from Senegal and Angola.
Family Chrysopetalidae
Dorsal surface of the segments bearing a fan or transverse row of paleae. Pros-
tomium with four eyes and three antennae. Biramous parapodia, with dorsal
cirri on all segments.
Genus BHAWANIA Schmarda, 1861
The head is exceedingly small and not easily seen ; the elongated body consists of
numerous segments. Neuropodial chaetae are compound.
Bhawania goodei Webster, 1884
Bhawania goodei, Augener, 1918, pp. 98-103, pi. 2, figs. 1-2 ; text-fig. 1 ; from Pram Pram.
Localities, (i) Accra, Christiansborg shore (1), almost complete, 8 mm. long.
(2) Axim, Hospital Reef, 7.i.5i (1), middle body piece, about 14 mm. long.
It has not been possible to examine the head in either of these specimens, since,
in the specimen from Accra, it has been damaged and it is missing from the Axim
example. The neuropodial heterogomph spinigers and falcigers are as figured by
Augener, but the paleae, which are of one type, are as shown in Text-fig. 3 and not
as illustrated by Augener. Monro (1933) noted that the tips of the paleae were not
emarginate.
Distribution. Apart from the above records B. goodei has been recorded from
Bermuda and False Bay, South Africa, in the Atlantic Ocean, and from the Galapagos
Islands and Taboga Island, Panama Bay, in the Pacific Ocean.
Family Amphinomidae
Body depressed, prostomium deeply set into the anterior segments, carrying three
antennae, two palpal pads and a caruncle. Biramous parapodia with branchiae
and simple chaetae (exceptionally uniramous with compound hooks).
Key to Genera
1. Body short and oval ; branchiae pinnate ....... Chloeia.
Body long and vermiform ; branchiae bushy . . . . . . . 2.
2. Branchiae beginning on the 1st chaetiger ; caruncle cushion-like, with dorsal surface
with chevron crenulations ......... Hermodice.
Branchiae beginning on the 2nd. or 3rd. chaetiger ; caruncle with a ventral crenulate
portion Eurythoe.
Genus CHLOEIA Savigny, 1818
Caruncle a plaited crest on a horizontal plate.
82
THE POLYCHAETE FAUNA OF THE GOLD COAST
Fig. 3. Bhawania goodei. One of the paleae from the dorsal surface.
(Specimen from Axim.)
Chloeia viridis Schmarda, 1861
Chloeia euglochis, Ehlers, 1887, pp. 18-24, pi- 1 > n S s - I_ ~ 2 > pi* 2 » n S s - I- 8 i pi- 3, n g s - I- 4-
Chloeia euglochis, Augener, 191 8, pp. 94-95, from Goree, Senegal.
Locality. Off Accra, Buchanan Survey, in 40 metres off Christiansborg (1),
70 mm. long by 10 mm. wide.
This is one of the few species of Polychaeta in which the pigment is retained, even
after long periods in preservative fluid. Thus, in this single specimen, the dorsal
cirri are violet, the pinnate gills, which appear at the fifth chae tiger, edged with
purple, whilst the superior neuropodial chaetae are a brilliant orange, as are a few
in the notopodial group. The remaining chaetae are glistening white, and the worm
presents a striking appearance. The caruncle is attached to the body to the second
chaetiger, though its unattached portion projects for 2-3 more segments, and it is
THE POLYCHAETE FAUNA OF THE GOLD COAST 83
surmounted throughout its length by a conspicuous, mid-dorsal, dark purple line.
Another thin mid-dorsal purple line, which is slightly broken in the intersegmental
furrows, extends along the back of the worm.
Distribution. This species is also known from the W. Indies, Gulf of California,
Galapagos and Cocos Islands.
Genus HERMODICE Kinberg, 1857
Caruncle cushion-like, with chevron crenulations on the dorsal surface.
Hermodice carunculata (Pallas) var. didymobranchiata (Baird), 1864
Amphinome didymobranchiata Baird, 1864, pp. 449-450, pi. 45, figs. 1-7 ; from Ascension
Island.
Hermodice carunculata var. didymobranchiata, Fauvel, 1914, pp. 113-116, pi. 8, figs. 22-27,
31-32 ; from the Gulf of Guinea.
Hermodice carunculata var. didymobranchiata, Monro, 1930, p. 27 ; from the Gulf of Guinea
and the French Congo.
Localities. Off Accra : Stn. 47 (3), 12, 20 and 30 mm. long ; Stn. 115 (6),
largest 40 mm. long, smallest 16 mm. ; Stn. 131 (1), 25 mm. long ; Stn. 133 (3),
2 complete, about 20 mm. long.
Most of these specimens have lost their colour except for the thin dark interseg-
mental bands on the back. The caruncle extends to the 4th chaetiger. Separation
of the branchiae into two bushy lobes instead of one and the almost complete absence
of harpoon-shaped dorsal chaetae, separates this variety from the stem species.
Distribution. This variety appears to have been recorded only from the above
listed areas.
Genus EURYTHOE Kinberg, 1857
Caruncle with vertical folds along the lateral walls, and with a vertical crenulate
portion.
Key to Species
1. Branchiae beginning on the second segment. Caruncle terminating on the third
or fourth segment. Four eyes ....... Ii. complanata.
2. Branchiae beginning on the third segment. Caruncle terminating on the first
segment. Four eyes ........ is. parvecarunculata.
Eurythoe complanata (Pallas), 1766
Eurythoe complanata, Augener, 191 8, pp. 88-89 ; from the Islands of Rolas and Annobon.
Eurythoe complanata, Fauvel, 1932, pp. 45-46.
Localities, (i) Tenpobo shore (70+). (2) Dixcove shore (7). (3) Axim,
(a) Hospital Reef (1), posterior fragment ; (b) Shore (1), small specimen.
These specimens vary considerably in length, the largest being about 100 mm. long,
and the smallest 15 mm.
Notopodial chaetae are of three types : (a) stout, straight and blunt ; (b) long and
calcareous, with a slender tip having a slight spur at the base ; (c) long harpoon-
84 THE POLYCHAETE FAUNA OF THE GOLD COAST
shaped, with lateral rows of teeth, and neuropodial chaetae of two forms : (a)
stout furcate chaetae with unequal arms ; (b) a few subfurcate with one arm thin and
elongated.
Distribution. This species is well known from littoral regions of the tropical
Atlantic, Pacific and Indian Oceans.
Eurythoe parvecarunculata Horst, 1912
Eurythoe parvecarunculata, Augener 1918, pp. 90-93, pi. 2, fig. 3 ; pi. 3, figs. 37, 38, from
Cameroons and Spanish Guinea.
Eurythoe parvecarunculata, Fauvel 19276, pp. 525-526, fig. 1 a-h, from Cameroons and French
Guinea.
Localities. Off Accra: Stn. 12 (1), anterior piece 17 mm. long; Stn. 16 (2),
anterior pieces 10 mm. and 15 mm. long ; Stn. 32 (3), two 20 mm. long, one 30 mm.
long ; Buchanan Survey, off Accra, in 3§-n metres, (1).
There are stout, harpoon-shaped, and slender, elongated, bifurcated chaetae in
the notopodial group. The bifurcation in the slender bristles may be reduced to a
spur. The neuropodial chaetae are furcate, slightly denticulate.
Distribution. This species is also known from the Indian Ocean, from Natal
to Malaya.
Family Phyllodocidae
Body normally long and slender with numerous segments. Prostomium variable
in outline, with two eyes and four or five antennae. Smooth or papillate proboscis,
protrusible and unarmed. Segments 1-3 bearing tentacular cirri. Parapodia
uniramous, with foliaceous dorsal cirri and compound chaetae.
Key to Genera
1. With 4 pairs of tentacular cirri .......... 2.
With 2 pairs of tentacular cirri ......... Eteone.
2. With 5 antennae ........... Eulalia. 1
With 4 antennae ........... Phyllodoce.
1 Not recorded from the Gold Coast.
Genus PHYLLODOCE Savigny, 1822
The four pairs of tentacular cirri are arranged on the three segments — 1 : 2 : 1.
Two anal cirri.
Key to Subgenera
1. Protruded proboscis proximally set with longitudinal rows of papillae . . (Anaitides).
2. Protruded proboscis proximally set with diffuse papillae . . . (Phyllodoce).
Key to Species of Anaitides
1. With more than 12 rows of papillae forming a continuous series around the proximal
half of the proboscis .......... A. africana.
2. With 12 rows of papillae arranged in two lateral groups of 6 on the proximal half of
the proboscis . , , , , A . oculata.
THE POLYCHAETE FAUNA OF THE GOLD COAST 85
Phyllodoce (Anaitides) africana Augener, 1918
Phyllodoce africana Augener, 1918, pp. 171-174, pi. 2, fig. 25 ; pi. 3, fig. 49-51 ; text-fig. 11.
Locality. Off Accra, Stn. 52, (1), incomplete, 24 mm. long.
The protruded proboscis is provided with a crown of six large bulbous papillae,
from which run six longitudinal rows of eight similar papillae. These rows are
followed by 24 longitudinal rows of about 14 much smaller papillae. Dorsal cirri
broadly foliaceous throughout, ventral cirri slender, those situated posteriorly
acquiring a pointed tip.
Distribution. This species has been previously recorded only from Angola
and Senegal.
Phyllodoce (Anaitides) oculata Ehlers, 1887
Phyllodoce {Anaitides) oculata, Augener, 1918, pp. 169-171, pi. 3, figs. 57-58, from Accra.
This species has not been found in the present survey.
Distally the proboscis is similar to the condition in A . africana, but the proximal
portion has only 12 rows of papillae, and these are disposed in two lateral groups of
six. Dorsal cirri become slender in the middle of the body.
Distribution. The W. Indies, Gulf of Guinea, Ascension Island and Tristan da
Cunha.
Genus ETEONE Savigny, 1822
Dorsal cirrus absent on the 2nd segment. Proboscis smooth, or with soft papillae
and small cuticularized tubercles.
Key to Subgenera
1. Proboscis smooth or with soft papillae ....... (Eteone).
2. Proboscis with lateral rows of large soft papillae and small spinous tubercles . (Mysta).
Eteone (Mysta) picta Quatrefages, 1865
Eteone {Mysta) picta, Fauvel, 1923, pp. 176-177, fig. 64, a-g.
Localities. Off Accra : Stn. 57 (1), 15 mm. long, incomplete ; Stn. 69 (2), 1
complete 17 mm., the other incomplete, 37 mm. long ; Buchanan Survey, 3-6 to
ii-o metres (4), 3 complete, 31, 38 and 42 mm. long.
For description these specimens may conveniently be divided into two groups :
one representing the young or immature form, the other the adult or sexually mature
condition.
(a) The immature forms (from Stns. 57 and 69). In these specimens the prostom-
ium bears two clear indentations on either side. The protruded proboscis is crowned
by an encircling group of large soft papillae, whilst two longitudinal rows of equally
large papillae extend laterally along the greater part of its length. Elsewhere there
are small papillae with soft chitinous teeth (Text-fig. 4, a). The dorsal surface of
the body in the specimen from Stn. 69 is marked with pale red-brown lateral stripes,
86
THE POLYCHAETE FAUNA OF THE GOLD COAST
although the effect of preservation may give the uniform stripes a speckled appear-
ance. In the specimen from Stn. 57 the colour is uniformly light brown, with dark
markings towards the sides. All specimens have the base of the foliaceous dorsal
cirrus coloured in pattern with the colouration of the adjacent dorsal surface of the
body. The remainder of the dorsal cirrus is greyish white or yellow, there being a
suture between these two clearly defined areas. Compound chaetae have long ter-
minal articles and project much further from the body than the cirri. On the articu-
Fig. 4. Mysta picta. (a) Small papilla, with chitinous teeth, from the proboscis, x 700
(b) compound chaeta, x 500. (Specimen from Stn. 69.)
lating surface of the main stem of the chaetae there are two teeth, as well as a small
finely pectinate plate (Text-fig. 4, b).
(b) The sexually mature forms (Buchanan Survey collections). In two of these
specimens the colour pattern is similar to that in the specimens described in group
(a), but the posterior parapodia contain small eggs, in which the nuclei are not visible.
The suture on the dorsal cirrus, however, is clearly seen and appears to segregate
a basal sac-like portion from a terminal solid piece. One of the remaining specimens
has much longer dorsal cirri extending posteriorly from the middle of the body.
In the anterior and middle regions of the dorsal surface the body is without colour
THE POLYCHAETE FAUNA OF THE GOLD COAST 87
but gradually becomes light bronze posteriorly. The lateral body-wall and the basal
sac-like portion of the parapodia, from behind the middle region of the body are rich
blood red, whilst the tips of the dorsal cirri are yellow. The basal portion contains
numerous large eggs, with visible nuclei ; the eggs are so large and numerous that
they produce a considerable swelling of the body- wall. The remaining specimen
has very much enlarged dorsal cirri from the 37th chaetiger, at which point the dorsal
body-wall and the bases of the parapodia have assumed a bronze colour, whilst the
tips of the dorsal cirri are yellow. Anterior to the 37th chaetiger the colour is similar
to that in specimens described in group (a). The parapodia, posteriorly from the
37th chaetiger, all contain eggs, the number to each segment increasing towards the
hinder end of the body, whilst the chaetae only project as far as the dorsal cirri.
Constant reference has been made to the prominent suture on the dorsal cirrus in
all the specimens here described, and examination of the mature specimens indicates
that the eggs collect in the basal sac-like portion of the parapodia, and it is here
suggested that the suture may represent the line of rupture when the eggs are liber-
ated through the body wall, into the sea, for external fertilization.
That these specimens represent a series from the immature to the sexually mature
condition is a supposition which can only be confirmed through observation on the
living animal. But the conclusion here reached — that all the specimens are of the
same species — appears justifiable, particularly when the variation in related genera
and families during their life histories is considered.
Distribution. This appears to be the first record of the species from the Equa-
torial Atlantic. Previously, E. (M.) picta has been reported from the North Sea,
the English Channel, the west coast of Scotland, the Bay of Biscay and the Mediter-
ranean.
Family Hesionidae
Body short, cylindrical. Prostomium simple or bilobed, normally with four eyes,
two or three antennae and two biarticulate palps. Protrusible proboscis with or
without jaws. Parapodia biramous, or reduced with long dorsal cirri and both
simple and compound chaetae.
Genus LEOC RATES Kinberg, 1865
Prostomium with four eyes, three antennae and two palps. There are eight pairs
of tentacular cirri. Proboscis with chitinous jaws. Parapodia biramous, noto-
podial chaetae simple, neuropodial chaetae compound.
Leocrates claparedii (Costa), 1868
Tyrrhena claparedii, Claparede, 1868, pp. 228-231, pi. 18, fig. 3.
Leocrates claparedii, Fauvel, 1923, pp. 237-238, fig. 88, i-n.
Locality. Axim, Hospital Reef, 13. iv. 49 (1), complete, 18 mm. long.
In this species the upper jaw plates have only one tooth, which distinguishes it
from L. atlanticus, with two teeth, reported by Fauvel (1950) from Dakar. The
median antenna in L. claparedii is short and subulate ; the paired laterals slender
88 THE POLYCHAETE FAUNA OF THE GOLD COAST
and slightly longer than the palps. Notopodial chaetae appear at the 5th chaetiger
and are simple and spinous : neuropodial chaetae have a bidentate sickle-shaped
terminal piece. There are 17 segments bearing parapodia, the last pair being
achaetous, retaining only the notopodial and neuropodial cirri as parapodial append-
ages, which, with the two urites, gives the appearance of there being six anal cirri.
Augener (1918) reported a new species, L. greeffianus, from the Isle of Rolas, by
St. Thomas, without noting the form of the pharyngeal armature.
Distribution. Originally recorded from Naples ; Day (1934 and 195 1) has
collected this species from the coast of Natal. It may have been found in the
Indo-Pacific region and reported under various names, but lack of details of the jaw
structure makes confirmation of this impossible.
Family Syllidae
Body small and narrow. Prostomium with four eyes, two palps and three anten-
nae. Normally two pairs of tentacular cirri on the first segment, which never has
chaetae. Protrusible proboscis, divided into two regions : (1) anterior, the pharynx,
chitinous, cylindrical, with or without teeth ; (2) posterior, the proventriculus, barrel-
shaped. Parapodia uniramous, normally with dorsal and ventral cirri. Chaetae
rarely simple, normally compound with a heterogomph articulation.
Key to Subfamilies
1. Without ventral cirri ......... Autolytinae. 1
With ventral cirri ............ 2.
2. Palps not fused ........... Syllinae.
Palps fused ............. 3.
3. Palps fused at the bases only ........ Eusyllinae.
Palps fused along their entire length ....... Exogoninae.
1 Not recorded from the Gold Coast.
Subfamily Syllinae
Key to Genera
1. A large tooth with a trepan ........ Trypanosyllis.
A large tooth with no trepan .......... 2.
2. With the tooth on the anterior part of the pharynx ..... Syllis.
With the tooth on the posterior part of the pharynx .... Opisthosyllis.
Genus TRYPANOSYLLIS Claparede, 1864
The proboscis is crowned with a circle of chitinous teeth (the trepan), and there is
also a large single anterior tooth.
Trypanosyllis prampramensis Augener, 1918
Trypanosyllis prampramensis Augener, 1918, pp. 276-278, pi. 4, figs. 91-92, text-fig. 26 ; from
Pram Pram.
Trypanosyllis prampramensis, Day, 1953, p. 414, from Still Bay and Kommetje, S. Africa.
This species is only known from these records, and has not been found in the present
survey. T. prampramensis appears to be very close to the European T. coeliaca,
THE POLYCHAETE FAUNA OF THE GOLD COAST 89
but is characterized by unidentate compound chaetae, the latter having bidentate
chaetae. Augener's specimen was 6-5 mm. long by 0-7 mm. wide for 100 segments.
Genus SYLLIS Savigny, 18 18
Prostomium with four eyes and often two anterior eye-spots. Pharynx crowned
with soft papillae and with a prominent anterior tooth. Antennae and dorsal cirri
are moniliform ; ventral cirri are not articulated.
Key to Subgenera
1. All chaetae simple .......... (Haplosyllis). 1
With compound chaetae and, sometimes, simple chaetae . . . . . 2.
2. Anterior chaetae compound, thereafter some large simple chaetae . . . (Syllis).
All chaetae compound . . . . . . . . . . . 3.
3. Compound chaetae with short articles all alike or differing only slightly from each
other ............ (Typosyllis).
Compound chaetae with long and short articles . . . . . (Ehlersia) .
1 Not recorded from the Gold Coast.
Syllis (Syllis) gracilis Grube, 1840
Syllis gracilis, Fauvel, 1923, p. 259, fig. 96, f-i.
Locality. Off Accra, dredge in 7-3 metres, 1 mile off shore, 2 miles west of
Densu River (1), 8 mm. long.
The dorsal cirri have between 8 and 12 articles. In the middle of the body chaetae
are simple (Text-fig. 5, a), but anteriorly and posteriorly there are compound falcigers.
The specimens here examined are not as long as those recorded by Fauvel (1923) ;
they may be immature.
Distribution. Mediterranean Sea, Atlantic Ocean, Red Sea, Indian and Pacific
Oceans.
Subgenus TYPOSYLLIS Langerhans, 1879
Key to Species
1. Dorsal cirri long with more than 20 articles ..... T. variegata.
2. Dorsal cirri short with less than 20 articles . . . . . . . T. hyalina.
Syllis (Typosyllis) variegata Grube, i860
Syllis {Typosyllis) variegata, Fauvel, 1923, p. 262, fig. 97, h-n.
Localities. Axim, (a) Shore, 13.iv.49 (1), 10 mm. long, incomplete; (b)
Hospital Reef, 7.i.5i (1), 7 mm. long, complete.
In the specimen from Axim shore, the dorsal cirri are alternately long and short,
with 30 to 40 and 20 to 25 articles respectively. The specimen from the Hospital
Reef, however, has dorsal cirri all about the same size with 30 to 40 articles. In
both specimens the pharynx is crowned with 10 to 11 soft papillae and has a promi-
nent anterior tooth. In the specimen from the Hospital Reef compound chaetae
go THE POLYCHAETE FAUNA OF THE GOLD COAST
have strongly bidentate, short, terminal articles, but in the specimen from the shore
the bidentation is less pronounced.
Distribution. English Channel, Mediterranean Sea, Atlantic Ocean, Indian
Ocean, Red Sea and Pacific Ocean.
Syllis (Typosyllis) hyalina Grube, 1863
Syllis (Typosyllis) hyalina, Augener, 1918, pp. 242-247, pi. 4, figs. 95, 96, from Pram Pram.
This species has not been collected in the present survey. It is distinguished
from T. variegata in having unidentate terminal articles on the compound chaetea,
and few articles on the dorsal cirri. T. hyalina frequently retains its colour after
preservation, showing a uniform light red anteriorly, or transverse bands of a red-
brown, but no specimens of T. variegata have been examined retaining any colouring
matter.
Distribution. English Channel, Mediterranean Sea and Atlantic Ocean.
Subgenus Ehlersia Langerhans, 1879
Syllis (Ehlersia) cornuta Rathke, 1843
Syllis (Ehlersia) sexoculata, Augener, 191 8, pp. 269-271, from Pram Pram in 9 metres depth.
Localities, (i) Off Accra, dredge in 7-3 metres, 1 mile off shore, 2 miles west
of Densu River, (1) , 6 mm. long. (2) Axim shore, 13 /14 . iv . 49, (2) , 5 and 6 mm. long.
There are two pairs of small eyes posterior to the median antennae, and two
minute eye-spots on the anterior border of the prostomium. Anterior dorsal cirri
have 8 to 10 articles, median cirri 10 and posterior cirri 4 to 5. The two anal cirri
have 12 articles. Compound chaetae are of two kinds, one with a long terminal
article (Text-fig. 5, b) and the other short (Text-fig. 5, c).
These specimens are small and may be juveniles.
Distribution. English Channel, Mediterranean Sea, Atlantic — Canaries, Senegal,
Angola — Persian Gulf and Indian Ocean.
Genus OPISTHOSYLLIS Langerhans, 1879
An occipital flap on the posterior border of the prostomium.
Opisthosyllis brunnea Langerhans, 1879
Opisthosyllis brunnea, Augener, 1918, pp. 274-276, text-fig. 25.
Localities, (i) Tenpobo shore, 3/4.U.50 (5), 3 complete, 8-10 mm. long.
(2) Winneba shore, 22.xi.49 (1), complete, 13 mm. long. (3) Axim, Hospital
Reef, 7.L51 (1), incomplete, 6 mm. long.
There are between 30 to 50 articles in the dorsal cirri. The pharynx extends over
eleven segments, bears a crown of 9 to n papillae and carries a large conical tooth
THE POLYCHAETE FAUNA OF THE GOLD COAST
9i
iJ\
Co)
Fig. 5. Syllidae chaetae. (a) Syllis gracilis, simple chaeta, x 300. (b) Ehlersia
cornuta, compound spiniger, x 400. (c) Ehlersia cornuta, compound falciger, x 400.
(d) Opisthosyllis brunnea, compound falciger, x 250. (e) Opisthosyllis brunnea,
simple chaetae, x 300.
posteriorly. The tooth is without colour and stands out against the dark brown of the
pharynx. Compound chaetae have short unidentate articles (Text-fig. 5, d). In
posterior parapodia there are long simple chaetae (Text-fig. 5, e).
Distribution. The tropical coast of W. Africa and the Indian Ocean.
Subfamily Eusyllinae
Genus PIONOSYLLIS Malmgren, 1867
Pharynx with a single anterior tooth.
ZOOL. 3, 2.
9 2 THE POLYCHAETE FAUNA OF THE GOLD COAST
Pionosyllis ehlersiaeformis Augener, 1913
Pionosyllis ehlersiaeformis, Augener, 1918, pp. 281-283.
Pionosyllis ehlersiaeformis, Day, 1953, pp. 415-416, text-fig. 3, d.
Locality. Off Accra, Buchanan Survey, in 14-5 metres off the Castle (1), in-
complete, 15 mm. long.
The pharynx is crowned with eight large papillae. Antennae and dorsal cirri
are very long with numerous articulations, which is unusual in this genus where,
normally, the appendages are smooth. Ventral cirri are smooth and pointed.
Superior chaetae have slender, curved unidentate terminal articles, about ten times
as long as the articles on the inferior chaetae which are straight with bidentate tips.
No eyes have been seen in this specimen, otherwise it is typical of the species.
Distribution. Augener originally recorded this species from S.W. Australia, but
in 1918 he reported it from German S.W. Africa, and in 1953 Day collected it from
Cape Agulhas.
Subfamily Exogoninae
Genus EXOGONE Oersted, 1845
Palps well developed, completely fused ; three antennae ; one pair of tentacular
cirri. Pharynx with a single tooth. Dorsal and ventral cirri small. Chaetae
simple and compound.
Exogone getnmifera (Pagenstecher)
Exogone gemmifera, Augener, 19 18, pp. 299-301, text-fig. 29, from Pram Pram in 9 metres
depth.
This species has not been collected in the present survey. Median antenna
longer than the prostomium and longer than the paired laterals ; two pairs of large
eyes. Dorsal cirri missing on the second chaetiger ; ventral cirri small, often difficult
to see. Pharynx straight, crowned with papillae, with a single anterior tooth.
In each parapodium there is a simple, bluntly terminated chaeta and a compound
form with a very small bidentate article ; on posterior feet there is a simple ventral
chaeta. E. gemmifera normally measures between 2-4 mm. in length.
Distribution. English Channel, Mediterranean Sea, and Atlantic Ocean.
Family Nereidae
Prostomium with four eyes, two antennae, two massive palps and four pairs of
tentacular cirri. Proboscis armed with two horny jaws and normally a series of
horny teeth (paragnaths). Parapodia normally biramous after the 2nd foot ;
chaetae compound ; most species have a Heteronereis, sexually mature form.
Generic and specific distinctions in this family are based primarily on the form
and arrangement of the paragnaths. These are divided first into two rings, the
maxillary and oral (Text-fig. 6), and then into eight numbered areas, of which I to IV
THE POLYCHAETE FAUNA OF THE GOLD COAST
93
are maxillary and V to VIII oral ; odd numbers are median and single, even numbers
lateral and paired. The jaws are at the distal end of the proboscis. The different
types of chaetae referred to in the text are illustrated in Text-figs. 8 and 12.
MAXILLARY RING
ORAL RING
BUCCAL SEGMENT
(a) W
Fig. 6. Diagrammatic representation of the division of the proboscis in Nereidae.
Key to Genera
1. Horny paragnaths of one type only, conical .....
Horny paragnaths of more than one type ......
2. Horny paragnaths of two types, conical teeth and transverse cutting plates
Horny paragnaths of three types, conical, transverse and pectinate
Nereis.
2.
. Perinereis.
Pseudonereis.
Genus NEREIS Linnaeus, 1758
All groups of paragnaths complete — Neanthes, sub-genus.
One or more groups of paragnaths missing — Nereis, sensu stricta.
Nereis falsa Quatrefages, 1865
Nereis callaona, Augener, 19 18, pp. 184-186.
Nereis falsa, Fauvel, 1923, pp. 337-338, rig. 129, e-m.
Nereis falsa, Day, 1951, pp. 27-28.
Localities, (i) Tenpobo shore (3), anterior pieces only, n-15 mm. long. (2)
(a) Accra, Christiansborg shore, 17. hi. 49 (20 -f), many complete, between 30-60
mm. long ; (b) Accra, on sponge near Sound, 9.xii.5o (1), anterior fragment 10 mm.
long ; (c) from wood netted off Chorkor, 14. v. 51 (1), complete, 18 mm. long. (3)
(a) Winneba shore (1), anterior fragment, 10 mm. long ; (b) Winneba, neap tide,
15.xi.49 (3), 20-30 mm. long. (4) Axim, (a) Shore 13/14.^.49 (4), 2 complete,
50 mm. long, others anterior fragments, 10-15 mm - l° n g '> (&) Hospital Reef, 7.L51
94
THE POLYCHAETE FAUNA OF THE GOLD COAST
(9), some complete, 10-15 mm - l° n g > i c ) Lighthouse Reef, 8.L51 (2), complete 10
and 20 mm. long.
This is one of the commonest littoral forms of the area. The arrangement of the
paragnaths varies very little from that shown in Text-fig. 7, though paired groups
VI sometimes have three or five teeth instead of two sets of four ; area V never has
any paragnaths. Notopodial chaetae are homogomph spinigers throughout, except
0-9
mm-
Fig. 7. Nereis falsa. Arrangement of the paragnaths on the proboscis,
(a) dorsal ; (b) ventral. (Specimen from Tenpobo.)
that in some of the posterior feet a single homogomph falciger may be present
(Text-fig. 8, a). Neuropodia have homogomph spinigers and heterogomph falcigers
dorsally and heterogomph spinigers (Text-fig. 8, b) and falcigers ventrally.
Distribution. — Atlantic, Mediterranean, Adriatic and around the South African
coast from Cape Town to Natal.
Nereis (Neanthes) succinea (Frey & Leuckart), 1847
Nereis glandulosa, Augener, 1918, pp. 192-194.
Nereis succinea Fauvel, 1936, pp. 307-313, {nee Nereis succinea Fauvel, 1923, pp. 346-347.)
Localities. Off Accra : (a) From wood netted of Chorkor, in 14 metres, 14. v. 51
(3), complete, 11-25 mm - l° n g '> (&) Stn. 29 (1), complete, 25 mm. long ; (c) 12 metres
off Accra (1), 38 mm. almost complete ; (d) Buchanan Survey, in a depth of 3-6 to
ii- o metres (10), between 25-60 mm. long, one a heteronereis.
The pattern of paragnaths varies considerably in this species around a typical
THE POLYCHAETE FAUNA OF THE GOLD COAST
95
form shown in Text-fig. 9. Thus I =1-4 in a longitudinal line ; II = arcs in two
rather scattered rows ; III = a transverse group in 2-3 scattered rows ; IV = arced
masses ; V = 0-4 (rarely 5-7). The absence of teeth on V in four of the specimens
from the Buchanan Survey does not necessarily upset the validity of the above key,
since this may be due to immaturity, accidental loss, general wear and tear, or it
may indicate the approach of sexual maturity. This latter reason is of doubtful
significance, but is referred to because this group is missing in the heteronereis
Fig. 8. Nereis falsa, (a) Homogomph falciger from a posterior notopodium, (b) hetero-
gomph spiniger from a neuropodium. Both X 250. (Specimen from Axim.)
example. Area VI is typically described as having a circle of six to seven paragnaths
surrounding a central one, but the majority of specimens here examined conform
to the pattern shown in Text-fig. 9, in which between 9-13 teeth are arranged in
three scattered rows. A specimen in the B.M. (N.H.), Reg. No. 1928.4.26.290,
from French Guinea, identified by Professor Fauvel, has 9-10 teeth in three scattered
rows, and this variation may be common among W. African examples of N. succinea.
Teeth in areas VII and VIII are both small and large and arranged, approximately,
in two rows.
A parapodium from the posterior region of one specimen is shown in Text-fig. 10.
The enlargement of the dorsal lobe only takes place posteriorly, anteriorly it is
long and thin. Notopodial chaetae are homogomph spinigers, neuropodial chaetae
96
THE POLYCHAETE FAUNA OF THE GOLD COAST
Fig. g. Nereis succinea. Arrangement of the paragnaths on the proboscis, (a)
dorsal, (b) ventral. (Specimen from the Buchanan Survey.)
THE POLYCHAETE FAUNA OF THE GOLD COAST
97
homogomph spinigers, and heterogomph spinigers and falcigers. The absence of
notopodial homogomph falcigers is considered by many workers (Fauvel, 1936 ;
Hartman, 195 1) to be important diagnostically. It is doubtful if a negative charac-
ter of this type can be so considered, since accidental loss of chaetae frequently occurs.
Distribution. Cosmopolitan ; frequently found in waters of low salinity.
0.5 mm
Fig. 10.
Nereis succinea. Posterior view of parapodium from the 32 nd chaetiger.
(Specimen from the Buchanan Survey.)
Genus PERINEREIS Kinberg, 1865
With transverse chitinous plates (horny paragnaths) on area VI ; other areas with
conical paragnaths.
Perinereis melanocephala Mcintosh, 1885
Nereis (Perinereis) melanocephala, Mcintosh, 1885, pp. 216-219, pi. 34 figs. 14-17 ; pi. i6a,
figs. 8, 9 ; from Bermuda, between tide marks.
Nereis (Perinereis) melanocephala, Augener, 1918, pp. 209-212, from the Island of Annobon and
the Cameroons.
Localities, (i) Tenpobo shore (8), 6 complete, 20-30 mm. long, others ant.
fragments : collected February, 1950. (2) Accra, Christiansborg shore (1), ant.
piece 38 mm. long. (3) Winneba shore, 15.xi.49, Neap tides (2), anterior pieces,
17-20 mm. long — one a developing heteroneis stage. (4) Sekondi, high level plateau,
21.ii.49 (3) complete, 20-25 mm. long. (5) Dixcove shore (1), complete, 64 mm.
long. (6) Axim, 13/14.1V.49 (1), complete, 18 mm. long.
9 8
THE POLYCHAETE FAUNA OF THE GOLD COAST
The proboscis of this species is shown in Text-fig. II. Little variation from the
normal has been found ; I = a triangular area of teeth ; II = rhomboidal area of
similar size to I ; III = rectangular area also equal to I ; IV similar to II ; V =
a single large tooth ; VI = single long, transverse, horny bands ; VII and VIII =
2 rows of teeth with scattered isolated teeth here and there. This compares accur-
ately with Mcintosh's original description. Unfortunately the type specimen,
B.M. (N.H.). Reg. No. 1885. 12. 1. 159 has no head. Efforts to find this portion,
including enquiries at the University of St. Andrews, where Mcintosh did his work,
have been unsuccessful. Notopodial chaetae are all homogomph spinigers (Text-fig.
mm.
Fig. 11. Perinereis melanocephala. Arrangement of the paragnaths on the proboscis,
(a) dorsal, (b) ventral. (Specimen from Sekondi.)
12, b), as are the superior neuropodial group. Inferior neuropodial chaetae are hetero-
gomph falcigers (Text-fig. 12, a). The superior notopodial lobe becomes enlarged
in the region of the 37th foot, continuing thus to the end of the body, the dorsal
cirrus being attached (Text-fig. 13, a) to its indented extremity. The degree of this
enlargement may be noted with respect to an anterior foot illustrated in Text-fig.
I3>&-
One of the specimens from Winneba has modified parapodia, beginning at the 17th
segment, indicating the development of the heteronereis stage (Text-fig. 14). The
lobe at the base of the dorsal cirrus, the post-chaetal neuropodial lobe and the ventral
cirrus are all in the early stages of development into foliaceous swimming surfaces.
The chaetae, however, remain normal, not having developed into the natatory type
expected in this genus, otherwise the aspect of the parapodia is similar to that
assumed by the more common P. cultrifera in the heteronereis condition.
Distribution, P, melanocephala is only known from the above records.
THE POLYCHAETE FAUNA OF THE GOLD COAST
99
Fig. 12. Perinereis melanocephala. (a) Neuropodial heterogomph falciger, x 500 ; (b)
notopodial homogomph spiniger, x 250. (Specimen from Sekondi.)
Genus PSEUDONEREIS Kinberg, 1865
With transverse chitinous plates (horny paragnaths) on area VI ; transverse rows
of pectinate teeth and conical paragnaths on other areas.
Pseudonereis gallapagensis Kinberg, 1865
Pseudonereis gallapagensis, Gravier, 1909, pp. 629-633, figs.
Pseudonereis variegata, Fauvel, 1927b, pp. 527-528, (part).
Pseudonereis gallapagensis, Hartman, 1948, pp. 68-69.
Localities, (i) (a) Winneba shore, 3 . iii . 50 (1), complete, 9 mm. long, 4.^.50(2),
one complete, 15 mm.; (b) Winneba rock face, 22.xi.49 (6), several complete,
THE POLYCHAETE FAUNA OF THE GOLD COAST
osrnm
Fig. 13. Perinereis melanocephala. Parapodia of (a) 37th chaetiger (specimen from
Sekondi) ; (b) an anterior chaetiger (specimen from Tenpobo).
OS
mm.
Fig. 14. Perinereis melanocephala. Parapodium from the 23rd chaetiger of a developing
heteronereid. (Specimen from Winneba.)
THE POLYCHAETE FAUNA OF THE GOLD COAST 101
between 15-30 mm. long; (2) (a) Axim shore, 13/14.1V.49 (1) complete, 19 mm.
long ; (b) Axim, hospital reef (2), both complete, one approaching sexual maturity,
17 mm. long, the other n mm. long.
Notopodial chaetae are homogomph spinigers; superior neuropodial chaetae
also homogomph spinigers, whilst the inferior group are heterogomph falcigers.
Posteriorly the dorsal lobe becomes foliaceous and carries at its distal end the dorsal
cirrus, leaving little or no free margin. Hartman (1948) referred to this character
as the specific distinction between P. gallapagensis and P. variegata (Grube), noting
that the latter species had a free portion at the tip of the dorsal foliaceous lobe.
Day (195 1) disputed the value of this distinction and it is doubtful if it is of practical
use. It is here suggested that P. gallapagensis has on each area IV of the proboscis
an apical group of about 16 small conical teeth in addition to the rows of pectinae
(Text-fig. 15, c), and this is a diagnostic character separating it from P. variegata.
Kinberg (1865) did not note this character, though his figures were not drawn in a
position which would show it. In examining the type specimen, which was in poor
condition, Hartman (1948) did not refer to the proboscidean structures. Gravier
(1909), however, observed this apical group on IV in specimens collected from Peru.
In groups VII and VIII in P. gallapagensis there is one row of alternating long and
short teeth (Text-fig. 15, b), the anterior tip of the long teeth being in line with the
Fig. 15. Pseudonereis gallapagensis. Arrangement of the paragnaths on the proboscis,
(a) dorsal, (b) ventral, (c) apical (jaws left out). (Specimen from Axim.)
latter. This is the condition on an extruded proboscis. If not extruded, however,
due to the different lengths of alternate teeth, and the general contracted condition
of the proboscis, it appears as if two rows of teeth are present. This character is
clearly shown in Kinberg's (1910) type figure and contrasts with the condition in
P, variegata in which there are 2/3 rows of short teeth (Mcintosh, 1903). All speci-
102 THE POLYCHAETE FAUNA OF THE GOLD COAST
mens here listed agree with this description, as does one from the Cameroons in the
B.M. (N.H.), Reg. No. 1928.4.2.844, which was recorded by Fauvel, 19276, as
P. variegata. Further specimens of P. gallapagensis in the B.M. (N.H.) collections
come from Ceylon and from Mormugas Bay, Goa, near Bombay.
Distribution. This is the first record of this species from the W. African coast ;
apart from the above records, P. gallapagensis is known widely from the western
coast of S. America, the Galapagos Islands and Hawaii.
Family Nephthydidae
Body with numerous short segments. Prostomium small and flat carrying two
pairs of antennae. Proboscis terminated with 14 to 22 soft bifid papillae and bearing
14 to 22 longitudinal rows of similar processes externally and two horny jaw pieces
on its inner wall. Parapodia are biramous except for the first pair which may not
be fully developed ; rami are wide apart with a coiled cirrus, probably branchiate
in function, between them.
Only three genera are recognized in this family and the following key has been
adapted from Hartman (1950) in which work an acceptable division of the type
genus Nephthys was suggested :
1. Inter-ramal cirri (branchiae) recurved — evolute ...... Nephthys.
2. ,, „ ,, involute ....... Aglaophamus.
3. ,, ,, ,, absent ....... Micronephthys.
No examples of the genus Micronephthys Freidrich, 1939, have been collected
from the Gold Coast, but Augener (1918) recorded M. ambrizettana from Angola.
Genus NEPHTHYS Cuvier, 1817
Nephthys hombergii Audouin & Milne-Edwards, 1830
Nephthys hombergii, Fauvel, 1923, p. 367, fig. 143, a-d.
Localities. Off Accra : Stn. 71 (1) 12 mm. long, anterior piece ; Stn. 132 (1)
16 mm. long, anterior piece ; Buchanan Survey, in 3-5 to 11 metres (2), one complete,
54 mm. long.
This well-known species is characterized by 22 longitudinal rows of papillae on
the proboscis, the appearance of the branchiae on the 4th chaetiger and the presence
of a notopodial button ventral to the acicula and tending to overgrow it.
Distribution. North Sea, English Channel, Mediterranean, Atlantic.
Genus AGLAOPHAMUS Kinberg, 1865
Aglaophamus lyrochaetus (Fauvel), 1902
Nephthys lyrochaeta Fauvel, 1902, pp. 72-5, figs. 9-12, from the estuary of the R. Casamance,
Senegal.
Nephthys lyrochaeta, Augener, 1918, pp. 160-166, pi. 2, fig. 12 ; pi. 3, fig. 59, from Saltpond,
Gold Coast in 9 metres, and Accra ; also several other W. African stations,
THE POLYCHAETE FAUNA OF THE GOLD COAST 103
Localities. Off Accra: Stn. 32 (1), anterior piece, 10 mm. long; Stn. 59 (1),
anterior piece, 7 mm. long ; Stn. 70 (2), both anterior pieces, 7 and 8 mm. long ;
Buchanan Survey, in 3-5-11 metres depth (4), two complete, 26 and 28 mm. long.
There are 14 longitudinal rows of papillae on the proboscis in this species. The
" lyre " bristles occur in the centre of each group of chaetae and are about half the
length of the others. They are therefore difficult to see unless the parapodium
chosen for observation is carefully mounted. A later diagnosis by Fauvel (19276) of
A . lyrochaetus indicated the presence of a small lobe on the dorsum of the neuropo-
dium which was confirmed by Monro (1930). With the " lyre " bristles, this character
serves as specifically diagnostic. The anterior feet are biramous and not uniramous
as indicated by Augener (1918), and the long ventral and dorsal cirri of the first
and second chaetigers respectively, noted by Monro (1930), are not present in the
specimens here described.
Distribution. — A . lyrochaetus is only known through the above records.
Family Glyceridae
Body elongate, tapering to both extremities, segments bi- or triannulate. Prosto-
mium conical, annulated, with four small terminal antennae. Protrusible proboscis,
very long, covered with papillae and armed with horny jaws of various kinds.
Normally with biramous parapodia, but frequently uniramous anteriorly. Branchiae,
when present, compound or simple, often retractile. Chaetae simple or compound.
Key to Subfamilies
1. Body divided into 2 or 3 regions by variation in the form of the parapodia which
may be uniramous, sub-biramous or biramous. Jaws and paragnaths numerous
Goniadinae.
2. Body not divided into distinct regions, parapodia either uniramous or biramous
throughout. Jaws and paragnaths few in number .... Glycerinae.
Subfamily Glycerinae
Key to Genera
1. Parapodia uniramous throughout, with compound cheatae only . . Hemipodus. 1
2. Parapodia biramous throughout, with simple chaetae in notopodia and compound
chaetae in neuropodia .......... Glycera.
1 Not recorded from the Gold Coast.
Genus GLYCERA Savigny, 1818
Prostomium long, with more than three annulations. Proboscis with four large
horny paragnaths distally and covered with papillae. Parapodia with a stumpy
dorsal cirrus, two anterior lobes one or two posterior lobes and a ventral cirrus.
Neuropodial chaetae compound spinigers, notopodial chaetae simple capillaries.
Branchiae present or absent, simple or branched, permanent or retractile into the
foot.
104
THE POLYCHAETE FAUNA OF THE GOLD COAST
Glycera convoluta Keferstein, 1862
Glycera africana, Arwidsson, 1898, pp. 21-22, pi. 1, figs. 10-12.
Glycera africana, Augener, 1918, pp. 384-386.
Glycera tridactyla, Augener, 1918, pp. 386-389, pi. 5, figs. 142-143, text-fig. 47.
Glycera convoluta, Fauvel, 1923, pp. 383-385, fig. 150, a-h.
Localities, (i) Off Accra: Stn. 65 (1); Stn. 69 (1) ; Stn. 89 (1) ; Stn. 126 (1) ;
all incomplete except Stn. 65, which measures 55 mm. long ; Buchanan Survey in
3- 6 to ii- o metres depth (6), five complete, 40-52 mm. in length. (2) Apam shore,
3/4.^.50 (3), one complete, 46 mm. long.
The papillae of this species are illustrated in Text-fig. 16, a and b. They are
cylindrical with a truncated termination, the truncation bearing a plate similar in
(c)
Fig. 16. Glycera convoluta. (a) Papilla from the proboscis, en face view, x 1500 (speci-
men from Apam) ; (b) papilla from the proboscis, side view, x 1500 (specimen from
Stn. 89) ; (c) silhouette of one of the 4 jaw articles, drawn in situ, (specimen from
Stn. 126).
appearance to the nail on the human finger. Over the greater part of the body both
anterior parapodial lobes are bluntly pointed and each project the same distance
from the body ; posterior lobes are unequal, the dorsal being blunt to oval and
projecting a short distance, whilst the ventral lobe is blunt and hardly projects at
all. Branchiae are non-retractile and lie on the dorsal edge of the parapodia. In
the anterior region they project as far as the chaetae, but posteriorly become very
much longer. Normally the branchiae appear at the I4th-i8th parapodia, but they
may not appear until the 29th foot. They are absent from extreme posterior segments.
A silhouette of one of the paragnaths is shown in Text-fig. 16, c, it was drawn in
situ, at the end of the protruded proboscis.
Distribution. Mediterranean Sea and Atlantic Ocean.
THE POLYCHAETE FAUNA OF THE GOLD COAST 105
Subfamily Goniadinae
Key to Genera
1. Proboscis with chevrons .......... Goniada.
Proboscis without chevrons . . . . . . . . . . 2 .
2. Neuropodia with spinigerous chaetae only ...... Ophioglycera,
Neuropodia with spinigerous and falcigerous chaetae .... Goniadopsis,
Genus GONIADA Audouin & Milne-Edwards, 1833
Body divided normally into two regions, an anterior uniramous and a posterior
biramous portion ; there may be a transitional region with sub-biramous parapodia.
Paragnaths include a pair of large dentate macrognaths separated from each other
by dorsal and ventral arcs of micrognaths. The proboscis carries two rows of dark
hard V shaped pieces, chevrons, and numerous papillae of one or few kinds, dis-
tributed in longitudinal, or irregular bands.
Goniada multidentata Arwidsson, 1899
Goniada multidentata Arwidsson, 1898, pp. 45-47, figs. 40-42.
Locality. Off Accra, Buchanan Survey in 39-6 metres depth, off Lagoon (1),
complete, about 90 mm. long.
There are 36 anterior uniramous, 15 median sub-biramous, and 131 posterior
biramous, parapodia. The uniramous feet bear prominent thick ventral cirri and
slender pointed dorsal cirri. There are three ligules to each chaetigerous lobe, two
anterior, long and finger-shaped, and one posterior, short. The sub-biramous portion
is so qualified only because the notopodial lobes are not well developed and also
because the segments in this region are the same width as those in the anterior
uniramous region. Notopodial chaetae, however, are developed and the segments
are strictly biramous. In the biramous posterior portion the segments are much
wider than anteriorly. Neuropodial ligules are as in the anterior region except that
the posterior ligule is triangular ; there are two notopodial ligules, both blunt, one
dorsal and the other ventral to the chaetae. Notopodial chaetae are acicular.
Neuropodial chaetae and chaetae in the anterior uniramous part are long hetero-
gomph spinigers (Text-fig. 17, a).
The large macrognaths each carry 12 teeth, six large and six small. There are 30
dorsal and 12 ventral micrognaths. Papillae from the proboscis are illustrated in
Text-fig. 17, b. About 150 pairs of chevrons have been counted, extending in two
rows along almost the full length of the proboscis. Arwidsson reported 90 pairs in
his original description, but it is not considered that the extra number present in
the Accra specimen is sufficient to warrant specific distinction. In all other characters
the specimens appear to be identical, and since the Accra specimen is almost twice
as long as Arwidsson's the variation may be taken to indicate differences in growth
stages.
Distribution. This species is known only from the tropical coast of West Africa.
io6
THE POLYCHAETE FAUNA OF THE GOLD COAST
Fig. 17. Goniada multidentata. (a) Chaeta from the anterior uniramous region ; (b)
group of papillae from the proboscis.
Genus GONIADOPSIS Fauvel, 1928
Body divided into three distinct regions ; (1) anterior, with uniramous parapodia
and stout falcigerous chaetae ; (2) median, with uniramous parapodia and spinigerous
chaetae ; (3) posterior, with biramous parapodia, notopodial acicular chaetae and
long spinigerous neuropodial chaeta. Papillae on the proboscis are numerous, of one
or few kinds.
Goniadopsis incerta Fauvel, 1932
Goniada (Goniadopsis) incerta Fauvel, 1932, pp. 122-123, pi- 4» n g s - 1-10, from Akyab, Burma.
Goniadopsis incerta, Day, 1953, p. 430, from the estuary of the Zwartkops River, Cape Province,
South Africa.
Locality, (i) Apam shore, 4.1V.50 (1), 35 mm. long.
The anterior region consists of 33 segments. Parapodia have three ligules, two
anterior, finger shaped, of equal length, one posterior, broad and triangular. One
of the stout falcigerous chaetae is shown in Text-fig. 18, a. Ventral cirri are short
and blunt anteriorly, but elongate in the posterior segments of this region.
THE POLYCHAETE FAUNA OF THE GOLD COAST
107
The median region consists of 37 segments. Parapodial ligules are similar to those
in the anterior region. One of the compound spinigers is shown in Text-fig. 18, b.
Ventral cirri are very long, dorsal cirri are short.
The posterior biramous portion makes up the remainder of the body of in seg-
ments. There are two notopodial ligules, one posterior ; neuropodial ligules are
Fig. 18.
Goniadopsis incerta. (a) Falciger from the anterior region ;
from the median region.
(b) spiniger
similar to those in the anterior region except that the posterior one is drawn out to
a blunt point. Compound spinigers are as in the median region.
There is one pair of small eye-spots at the base of the annulated prostomium.
Distribution. This is the first record of this genus from the Atlantic Ocean.
Genus OPHIOGLYCERA Verrill, 1885
Body divided into three regions ; (1) anterior with uniramous parapodia, with
spinigerous chaetae ; (2) median, in which notopodia with simple chaetae are
gradually developed ; (3) posterior, with biramous parapodia. Proboscis long,
terminated with a circlet of soft fleshy papillae, within which is a circlet of macrog-
naths. Papillae on the proboscis are numerous of one or few kinds.
Ophioglycera archeri sp. n.
Holotype, B.M. (N.H.), Reg. No. 1953.3. 1. 648-
Locality. Off Accra, Buchanan Survey, in 7-3 metres off Castle (1), complete,
134 mm. long. The proboscis, which is not protruded, measures about 25 mm. long.
The prostomium is without visible annulations or eye-spots. Eighteen large
papillae form a crown at the end of the proboscis. The proboscideal teeth form a
ZOOL. 3, 2. 9
io8
THE POLYCHAETE FAUNA OF THE GOLD COAST
circlet, adjacent to the papillae, of two large separate macrognaths having between
them groups of 18 and 25 micrognaths. The macrognaths have four teeth each,
two large and two small, the micrognaths are bidentate. Papillae cover the surface
of the proboscis and appear to be all alike (Text-fig. 20, b).
On the first foot there is a single cirrus with no chaetae. Thereafter there are 27
uniramous parapodia with spinigerous chaetae (Text-fig. 19, a), 49 segments in
which notopodia are developed, but in which the rami are close together (Text-fig.
19, b) and 140 biramous parapodia in which the rami are far apart (Text-fig. 20, a).
Neuropodial chaetae are compound spinigers throughout, notopodial chaetae are
simple.
In this specimen biramous parapodia are not gradually developed. At the 28th
chae tiger the notopodial ramis appear fully developed with lobes and simple chaetae.
A third division of the body, however, is recognizable at the 77th chaetiger where
notopodia and neuropodia are much wider apart than they are anteriorly, and the
chaetigerous ligules are much more foliaceous (compare Text-fig. 19, a and 19, b
with Text-fig. 20, a).
No diagnostic coloration is apparent in the preserved condition of the worm.
No other species of Ophioglycera is known in which the change in form of the
parapodia begins as far forward as in 0. archeri. In 0. foliocea the change begins at
about the 35th chaetiger, but in 0. longicirrata, 0. gigantea, 0. eximia and 0. distorta
it does not appear until the 55th or 59th, and, exceptionally, may be delayed to
the 90th.
<9 (9
Fig. 19. Ophioglycera archeri, sp. n. Parapodia, (a) 13th foot, (b) 28th foot.
THE POLYCHAETE FAUNA OF THE GOLD COAST
109
■07
mm
Fig. 20. Ophioglyczva avchevi, sp. n. (a) Parapodium from a posterior foot of a mature
$ ; (b) papilla from the proboscis.
no THE POLYCHAETE FAUNA OF THE GOLD COAST
Family Eunicidae
Body elongated, with the prostomium normally having one pair of palps and one
to seven antennae. First two segments normally achaetous and apodous. Some-
times one pair of tentacular cirri on the 2nd segment. Parapodia uniramous or
sesquiramous. Dorsal cirrus, with or without branchiae, sometimes missing or
rudimentary ; ventral cirrus often absent. Chaetae simple and /or compound,
showing great variety of shape. Proboscis armed with a ventral mandible and dorsal
maxillae. A membranous tube is secreted by some species.
Key to Sub-families
i. Prostomium reduced, without appendages (except in Augenaria Monro, with three
minute antennae). No ventral cirri ; dorsal cirri rudimentary . . Lumbrinerinae.
Prostomium with appendages, palps and antennae . . . . . . 2.
2. No ventral cirri ; dorsal cirri foliaceous. Three antennae . . . Lysaretinae.
Ventral cirri present ............ 3.
3. Two antennae. Maxillae of two or four longitudinal series of very small and numerous
pieces ............ Dorvilleinae. 1
One to seven antennae. Maxillae of four or five pairs of pieces . . . . 4.
4. Seven antennae, five occipitals mounted on ringed ceratophores and two ovate
frontals ........... Onuphidinae.
One to five antennae, ovate frontals absent ...... Eunicinae.
Sub-family Eunicinae
Key to Genera
1. Branchiae present. Five antennae . . . . . . . . . 2.
Branchiae absent ............ J.
2. Tentacular cirri present .......... Eunice.
Tentacular cirri absent .......... Marphysa.
3. Three tentacles, tentacular cirri absent ....... Lysidice.
One tentacle, tentacular cirri present ....... Nematonereis. 1
Genus EUNICE Cuvier, 1817
Five antennae. One pair of tentacular cirri on the second apodous segment.
Dorsal cirri elongated, ventral cirri short or knob-like. Branchiae simple or pinnate.
Parapodia sesquiramous, with acicular chaetae. Simple pectinate (comb-like) chaetae,
and compound chaetae. Mandible of two pieces. Maxillae with a pair of forceps
and two or three pairs of toothed plates, an unpaired left plate and sometimes
paragnaths.
There are manifold difficulties in identifying incomplete specimens of Eunice. The
most important of these concerns the arrangement of those characters which appear
on every parapodium, thus necessitating complete specimens for accurate diagnosis.
A high degree of variability in the arrangement of branchiae, jaw plate dentition
and extent of antennal annulation within any one species introduces further un-
certainty. For these reasons all specific identifications of incomplete specimens are
doubtful.
1 Not recorded from the Gold Coast.
THE POLYCHAETE FAUNA OF THE GOLD COAST in
Key to Species
i. Branchiae normally absent ; when present, simple, posterior, with one filament,
sometimes 2-3 . . . . . . . . . . . E. gracilis.
Branchiae, always present, pinnate . . . . . . . . .2.
2. Subacicular hooks bidentate .......... 3.
Subacicular hooks tridentate .......... 4.
3. Branchiae occur only on the anterior part of the body beginning at about the 6th
chae tiger ........... E. coccinea.
Branchiae occur only on the middle and posterior part of the body, beginning at
about the 26th chaetiger ........ E. filamentosa.
4. Branchiae first present from about the 3rd chaetiger but absent over the greater part of
the body from the middle backwards ...... E. vittata
Branchiae first present from the 3rd to 7th chaetigers, and also present over the
greater part of the posterior region . . . . . . . . . 5.
5. Branchiae present on the last few chaetigers . . ■ . . E. antennata.
Branchiae absent from the last few chaetigers . . . . . E. rubra.
Eunice antennata Savigny, 1818
Eunice antennata, Crossland, 1904, pp. 312-318, figs. 56-60.
Eunice antennata, Hartman, 1944a, pp. 115-117, pi. 7, figs. 154-156.
Localities, (i) Tenpobo shore, (1), anterior fragment, 28 mm. long. (2) Off
Accra, Dredge in 14-6 m., shingle, 2 miles south out from the R. Densu (1), 70 mm.
long, complete, and several fragments.
The antennae are clearly annulated. Pectinate branchiae, in the dredged specimen,
begin on the 7th chaetiger with 9 filaments ; on the 24th, 32nd and 52nd feet there
are 12, 4 and 1 filament to each gill respectively. An increase to 3 filaments then
takes place, which is maintained up to the last three segments on all of which there
is a further reduction to one. Subacicular hooks are tridentate, compound chaetae,
bidentate falcigers ; simple chaetae are capillary and pectinate. There are two
pairs of anal cirri, one dorsal and long, the other ventral and short.
Distribution. Cosmopolitan in tropical and subtropical areas.
Eunice vittata (Delle Chiaje), 1829
Eunice vittata, Augener, 1918, pp. 321-323.
Eunice vittata, Fauvel, 1923, pp. 404-405, fig. 158, h-n.
Eunice vittata, Hartman, 1944a, p. 118.
Localities, (i) Tenpobo shore, (1), ant. piece. (2) Off Accra : (a) Stn. 35 (1),
complete, 14 mm. long ; Stn. 47 (2), anterior pieces ; Stn. 69 (1) anterior piece ;
Stn. 88 (1), complete, 8 mm. long; (b) Dredge in 14-6 m., shingle, 2 miles south out
from the R. Densu (2), complete, 10 and 19 mm., long, and nine anterior pieces.
In the specimen from Stn. 88 branchiae appear on the 3rd chaetiger with three
filaments. A maximum development of 7-8 filaments occurs on the 38th foot, after
which there is a reduction to one and finally branchiae are missing entirely from the
posterior segments. Subacicular hooks, and chaetae, are similar to those in E.
antennata.
Distribution. Mediterranean, Atlantic, and Pacific.
THE POLYCHAETE FAUNA OF THE GOLD COAST
Eunice ftlamentosa Grube, 1857
Eunice filamentosa, Augener, 1918, pp. 324-327, from Pram Pram.
Eunice filamentosa, Hartman, 1944a, p. 107, pi. 6, figs. 123-126.
Localities, (a) Axim shore, 13/14.^.49 (1), 40 mm. long, complete ; (b) Axim,
Hospital Reef, 7.L51 (1), 200 mm. long, complete.
In the specimen from Axim shore branchiae appear at the 29th chaetiger with one
filament, gradually increase to a maximum of five and continue thus over the greater
part of the body. Towards the pygidium there is a reduction to 3/4 filaments. The
subacicular hooks which appear at the 23rd foot are strongly bidentate ; superior
acicular have blunt, hammer-shaped ends.
The specimen from the Hospital Reef has branchiae appearing at the 26th chaetiger,
the maximum number of filaments is five and this condition prevails over much of
the body. There is again a gradual reduction in filament number towards the tail,
but in this specimen it is carried much further, the last three chaetigers having no
branchiae. Fragments of a papyraceous tube are present.
Distribution. Tropical West Africa, Eastern and Western Tropical America.
Eunice coccinea Grube, 1878
Eunice coccinea, Crossland, 1904, pp. 297-303, pi. 20, figs. 6, 7, text-figs. 46-51.
Eunice coccinea, Fauvel, 1932, p. 136.
Locality. Axim, Hospital Reef, 7.L51 (2), one complete, 42 mm., one incomplete,
100 mm. long.
Branchiae begin on the 6th and 7th parapodia. They attain a maximum of 10-12
filaments in the anterior third and are missing throughout the rest of the body.
Subacicular hooks appearing at the 25th and 28th feet, are bidentate.
Distribution. Frequently recorded in the tropical regions of the Atlantic and
Indo-Pacific, from the Gulf of Guinea to the Philippines.
Eunice gracilis (Crossland), 1904
Nicidion gracilis, Crossland, 1904, pp. 327-329, pi. 22, figs. 10-11, text-figs. 65-66.
Eunice gracilis, Fauvel, 1932, pp. 140-141, text-fig. 20.
Eunice gracilis, Day, 1949, p. 447.
Localities, (i) Accra, (a) Christiansborg shore, 17.iii.49 (2), complete, 7 and
n mm. long ; (b) Dredge 8 fathoms, shingle, 2 miles south of R. Densu (1), complete,
20 mm. long. (2) Dixcove, shore (10), five complete, 15-42 mm. long. (3) Axim, (a)
Shore, 13/14.^.49 (1), complete, 24 mm. long; (b) Lighthouse Reef, 8.L51 (1),
complete, 20 mm. long; (c) Hospital Reef, 13.iv.49 (1), complete, 22 mm. long;
7.L51 (10), seven complete, 10-18 mm. long.
Simple chaetae are capillaries and comb-like ; compound chaetae are bidendate.
Subacicular chaetae appear at about the 25th foot and are also bidendate ; there is
one to each parapodium compared with two in E. cincta Kinberg, (see Hartman,
THE POLYCHAETE FAUNA OF THE GOLD COAST 113
1948, p. 80, on a re-examination of Kinberg's types). For this reason the synonymy
proposed by Fauvel (1950) including E. gracilis as a synonym of E. cincta has not
been accepted here. Antennae barely reach back to the 1st chaetiger ; tentacular
cirri have frequently been lost. No branchiae are present ; anteriorly dorsal cirri
are well-developed.
There has been much discussion about the generic status of this species. Originally
designated a species of Nicidion because of the lack of branchiae, Fauvel (1932)
and Day (1949) have reported gills sometimes present on extreme posterior segments,
and for this reason the species is regarded as a Eunice.
Distribution. Indian Ocean — Zanzibar, Gulf of Manaar, Mergui peninsular —
and Atlantic Ocean. St. Helena and the Gold Coast.
Eunice rubra Grube, 1856
Eunice rubra, Augener, 1918, pp. 319-321, from Pram Pram, and the Island of Rolas, in the
Gulf of Guinea.
This species has not been found in the present collections. The antennae are
clearly annulated, branchiae begin at the 5th chaetiger and subacicular hooks are
trident ate.
Distribution. This species is common on the south-eastern coast of the U.S.A.
Genus MARPHYSA Quatrefages, 1865
Five antennae, two eyes. Tentacular cirri absent, dorsal cirri elongated ;
ventral cirri short. Branchiae simple or pectinate. Dorsal chaetae simple capillaries.
Ventral chaetae simple or compound, comb chaetae, and acicular chaetae also
present. Mandible of two pieces. Maxillae with a pair of forceps, two pairs of
toothed plates, an unpaired plate and sometimes paragnaths.
Marphysa dartevellei Monro, 1936
Marphysa dartevellei Monro, 1936&, pp. 246-248, 6 figs, from the Congo coast, Cape Malemba,
85 km. north of Banana, from holes of Lithodomus.
Localities, (i) Tenpobo shore, 5.U.50 (1), 325 mm. long, complete. (2) Accra,
Christiansborg shore, 19.xi.49 (2), anterior pieces, 85 and 35 mm. long. (3) Axim,
(a) Shore, 13/14.^.49 (2), anterior pieces, 26 and 35 mm. long ; (b) Hospital Reef,
7.1.51 (1), 58 mm. long, complete.
The antennae are unarticulated, equal in length, measuring only a little longer
than the head ; between the laterals there are two black eyes. Dorsal cirri are smooth
and slender decreasing in length posteriorly. Ventral glandular pads are present
after the first few chaetigers. Ventral cirri are well developed stumps in the anterior
feet, but posteriorly become little more than tubercles at the top of the glandular
pads. Pectinate branchiae begin between the 32nd to 40th chaetigers. They have
a maximum of four to five filaments and are present over the greater part of the
middle body region. In the complete specimen from Tenpobo there are 86
ii 4 THE POLYCHAETE FAUNA OF THE GOLD COAST
abranchiate segments posteriorly preceded by a large number of segments in which
reduction to one filament has taken place.
In the anterior parapodia compound chaetae are spinigers and posteriorly are
falcigers. Monro's original specimens were not complete and his suggestion that
falcigerous compound chaetae replace spinigers posteriorly is here substantiated.
Simple capillary chaetae occur throughout the body, simple comb-chaetae are present
in the middle body region but are often difficult to find posteriorly.
There are two pairs of anal cirri, one pair elongated and dorsal, the other ventral
and short.
Distribution. This species is known only from the above records.
Genus LYSIDICE Lamarck, 1818
Three antennae. Tentacular cirri absent. Branchiae absent. Chaetae, simple
capillaries, comb-like, compound falcigers and acicular. Mandible of two pieces.
Maxillae with a pair of forceps, two toothed plates, an unpaired plate and paragnaths.
Key to Species
1 . Eyes round ; antennae small, never reaching as far as the anterior border of the
prostomium ............ L. ninetta.
2. Eyes reniform ; antennae reaching to the anterior border of the prostomium, some-
times further . . . . . . . . . . . L. collaris.
Lysidice ninetta Audouin & Milne-Edwards, 1834
Lysidice ninetta, Augener, 191 8, pp. 362-364.
Lysidice ninetta, Fauvel, 1923, pp. 411-412, fig. 162, a-g.
Localities. Off Accra : Stn. 48 (1), 13 mm. long, complete ; Stn. 62 (1),
6 mm. long, anterior piece ; from wood netted in 14 metres off Chorkor (1),
21 mm. long, complete.
For further details see below under L. collaris.
Lysidice collaris, Grube, 1869
Lysidice collaris, Crossland, 1903, pp. 143-144.
Lysidice collaris, Fauvel, 1932, pp. 143-144.
Lysidice collaris, Day, 195 1, pp. 39-40.
Locality. Off Accra, Stn. 62 (1), 22 mm. long, complete.
Differences, other than those noted in the above key, which serve to separate
these two species are (a) a marked indentation on the anterior border of the prosto-
mium compared with the almost entire border in L. ninetta, and (b) the greater
development of the dorsal cirri in L. ninetta.
The form of the jaw articles do not differ sufficiently to warrant emphasis. Day
(195 1) has shown that this character is of value in differentiating between L. collaris
and L. natalensis, particularly with respect to maxillae II. In L. collaris and L.
THE POLYCHAETE FAUNA OF THE GOLD COAST 115
ninetta, however, the former has four teeth on each article of maxillae II, the latter
varying between 4-4 and 4-5.
Chaetae are similar in both species. The simple types, capillaries and comb-like,
are dorsal, the compound, with bidentate terminal articles, are ventral. In both
species there are two pairs of anal cirri, a long dorsal pair and a short ventral pair.
Distribution. L. ninetta is known extensively from, the European area in which
L. collar is is unknown. There appears, however, to be a general overlap of the two
species in the Pacific Panama, West African and Indian Ocean regions. L. collaris
has been frequently recorded from the Indo-Pacific region, from Australia, China,
Philippines, Japan, etc., in which areas no valid record of L. ninetta appears to have
been made. The record of this latter species from New Zealand (Fyfe, 1952, p. 19)
doubtfully accepted by Fauvel (1923, p. 411) and detailed by Augener (1924) as
L. brevicornis Kinberg, is almost certainly L. collaris.
Subfamily Onuphidinae
Key to Genera
1. Branchial filaments inserted spirally ........ Diopatra.
2. Branchial filaments simple or pectinate ....... Onuphis.
Genus DIOPATRA Audouin & Milne-Edwards, 1833
Seven antennae, of which two are small front als and five occipitals borne on ringed
ceratophores. One pair of clear, translucent, circular areas posterior to the inner
laterals, which are probably sensory in function. One pair of tentacular cirri on
the apodous peristomium. Dorsal cirri subulate ; ventral cirri subulate in a few
anterior segments thence becoming pad-like. Pseudo-compound chaetae in anterior
segments, otherwise with simple capillary, comb-like and acicular chaetae. Branchiae
large with filaments inserted spirally. Mandible of two pieces. Maxillae with a pair
of forceps, three pairs of toothed plates and one unpaired plate. Tube parchment-
like with agglutinated material of varying types.
Key to Species
1. Tip of the shaft of the comb-chaetae continued as a central spike (Text-fig. 21, a).
Tube as in Text-fig. 22, a . . . . . . . . D. musseraensis.
2. Tip of the shaft of the comb-chaetae not so continued. Tube as in Text-fig. 22, b.
D. neapolitana.
Diopatra musseraensis Augener, 1918
Diopatra musseraensis Augener, 1918, pp. 347-349, pi. 5, fig. 134 ; pi. 6, fig. 195 ; text-fig. 38,
from Angola.
Localities, (i) Off Accra, dredge, 14-6 metres, in shingle 2 miles south from the
mouth of the R. Densu (1), incomplete, 40 mm. long. (2) Winneba shore, 22.xi.49
(5), incomplete, 25-120 mm. long, with tubes measuring 160-240 mm. in length.
Strongly spiralled branchiae are present on about 50 chae tigers. The anterior
dorsal surface is uniformly coloured mauve.
n6
THE POLYCHAETE FAUNA OF THE GOLD COAST
Chaetae are in well-defined groups. Prominent comb-chaetae (Text-fig. 21, a) are
dorsal and anterior, they project almost as far as the stout winged capillaries which
form a large group ventral to them. A bulbous parapodial cirrus projects beneath
this latter group and divides the anterior set of pseudo-compound bristles from a
Fig. 21. Comb-chaetae of (a) Diopatra musseraensis , x 500 ; (b) and (c)
Diopatra neapolitana, x 750.
small posterior group of capillaries. The pseudo-compound chaetae number four
to a parapodium and have a darker brown terminal piece apparently grafted on to
the light brown basal portion. This gives the effect of a possible articulating surface.
Ventro-anteriorly to the ramal cirrus are two bidentate subacicular chaetae, which
first appear at about the 17th parapodium.
Distribution. Known only from the above records.
Diopatra neapolitana Delia Chiaje, 1841
Diopatra cuprea, Augener, 1918, pp. 350-354, text-fig. 39, from Elmina and Accra, Gold Coast,
etc.
Diopatra neapolitana, Fauvel, 1923, pp. 419-420, fig. 166, a-h.
Diopatra neapolitana, Fauvel, 1933, pp. 28-37, n g s - 3 _ 4-
Localities. Off Accra : Stn. 6 (1), + tube ; Stn. 16 (1) ; Stn. 27 (1), +
tubes ; Stn. 53 (1) ; Stn. 57 (34), -f tubes ; Stn. 58 (6), + tubes ; Stn. 59 (1) ;
Stn. 61 (1), -f tubes ; Stn. 70 (6) ; Stn. 73 (6), + tubes, one specimen complete,
30 mm. long ; Stn. no (1), with tube ; Stn. in (1), with tube ; Stn. 126 (1), com-
plete, 27 m. long. Apart from the specimens from Stns. 73 and 126, all are less
THE POLYCHAETE FAUNA OF THE GOLD COAST
117
than 15 mm. in length and are anterior fragments. Buchanan Survey, 3-6 to iro
metres off Accra (7), incomplete, up to 30 mm. in length.
In the complete specimen from Stn. 126 branchiae begin on the 5th parapodium
and are spiralled up to the 30th foot, after which they become branched with one
or two filaments for 10 segments and then disappear. The anterior dorsal surface
is skin-coloured, crossed, in some specimens, by lateral stripes of pale mauve.
mm
®
Fig. 22. Tubes of (a) Diopatra musseraensis, (b) Diopatra neapolitana.
The form of the comb-chaetae in these specimens is illustrated in Text-figs. 21, b-
c. These chaetae were exceedingly difficult to find and in no specimens were they
as liberally distributed among the parapodia as in D. musseraensis.
Fauvel (1932 & 1933) has drawn attention to the variation in form of the comb-
chaetae in this species, and examination of the Gold Coast specimens supports his
conclusions. In one specimen, for example, comb-chaetae with 6 and 14 teeth are
present. For this reason Augener's record of 1918 of D. cuprea has been included as
a synonym of D. neapolitana. With the exception of the scarcity of comb-chaetae
n8 THE POLYCHAETE FAUNA OF THE GOLD COAST
in D. neapolitana the distribution of chaetae in the parapodia is very similar to that
in D. musseraensis.
Distribution. Atlantic Ocean, Mediterranean Sea, Red Sea, Indian and Pacific
Oceans.
Genus ONUPHIS Audouin & Milne-Edwards, 1833
Similar to Diopatra except that branchiae are not spirally coiled, but either simply
branched or pectinate.
Onuphis eremita Audouin & Milne-Edwards, 1833
Onuphis landanaensis, Augener, 1918, pp. 339-343, pi. 5, figs. 135-138, pi. 7, fig. 197, text-
fig- 36.
Onuphis eremita, Fauvel, 1923, pp. 414-415, fig. 163, a-l.
Localities, (i) Tenpobo shore, about 30 specimens with tubes, some complete,
measuring 30 mm. in length ; associated with Audouinia punctata. (2) Off Accra :
Stn. 57 (3), fragments, up to 18 mm. long ; Stn. 12 (1), anterior piece.
Eyes are absent in this species. Branchiae begin on the first chaetiger with one
filament ; after 10 to 20 segments there is a gradual increase to a maximum develop-
ment of five filaments. There are tri- or bidentate hooded pseudo-compound chaetae
in the first three to five feet. Other chaetae are simple limbate capillaries and
comb-like ; acicular chaetae are bidentate. The specimens from Tenpobo are
much smaller than those described by Fauvel. Tubes are of agglutinated sand
grains with a few shell fragments.
Distribution. Atlantic Ocean, Mediterranean Sea, Suez Canal, Madagascar,
Ceylon, Madras, Mergui, Akyab, Caribbean Sea, and off the coast of South California
down to Guatemala.
Subfamily Lumbrinerinae
Genus LUMBRINERIS Blainville, 1828
Body long and cylindrical. Prostomium without palps, antennae or eyes. First
two segments apodous and achaetous. Branchiae absent. Parapodia with two
unequal ligules. Simple winged capillary chaetae and simple or compound hooks.
Mandible bodice-like. Maxillae with four pairs of plates and two supports.
Key to Species
1 . Prostomium conical, pointed anteriorly ; no compound hooded hooks, simple hooks
as Text-fig. 23, c . . . . . . . . . . L. impatiens.
2. Prostomium rounded, blunt anteriorly with some compound hooded hooks (Text-
fig. 23, a), simple hooks as in Text-fig. 23, b . . . . L. albifrons.
Lumbrineris impatiens (Claparede), 1868
Lumbriconereis impatiens, Fauvel, 1923, pp. 429-430, fig. 171, a-i.
Localities, (i) Tenpobo shore, 4.ii-5o (1), anterior piece 13 mm. long. (2) Accra,
Christiansborg shore (2), one complete, 78 mm. long ; one fragment, 20 mm. long.
THE POLYCHAETE FAUNA OF THE GOLD COAST
119
Parapodia with a short, anterior, rounded lobe, and a long, posterior, cirriform
ligule. Anterior parapodia have simple winged chaetae and simple hooks with
denticles and guard. The capillaries are gradually replaced in the middle and posterior
region, leaving only two or three simple hooks (Text-fig. 23, c) to each chae tiger.
The large maxillary plates II have four teeth each, III 2 +2 and IV 1 + 1. The
maxillary supports are much longer than those described by Fauvel.
Distribution. Atlantic Ocean, Mediterranean, Red Sea, Persian Gulf, Indian
Ocean.
Fig. 23.
Chaetae of Lumbrineris. (a) Compound hooded hook of L. albifvons, (b) simple
hooded hook of L. albifvons, (c) simple hooded hook of L. impatiens.
Lumbrineris alhifrons (Crossland), 1924
Lumbriconereis albifvons Crossland, 1924, pp. 50-55, text-figs. 65-72, from the Cape Verde
Islands.
Locality. Off Accra, from wood netted off Chorkor 24. v. 51 (2), complete, 75
and 22 mm. long.
This species is characterized by the form of the compound chaetae (Text-fig. 23, a).
These are, however, exceedingly difficult to find. The dental apparatus is illustrated
in Text-fig. 24 and also serves to distinguish the species from closely related forms.
Simple winged capillary chaetae are present with hooded hooks anteriorly, but are
replaced by the latter posteriorly. As in L. impatiens, anterior parapodial lobes are
short, posterior lobes are long.
Distribution. The only authentic records of this species are those noted above ;
the records by Monro (1933) from the Galapagos Islands and Pruvot (1930) from,
New Caledonia are of doubtful significance.
Subfamily Lysaretinae
Apart from the characters noted above in the key, the following are of importance
in this subfamily. Mandible of two pieces, maxillae with five pairs of symmetrical
toothed plates and two long supports (Text-fig. 25).
THE POLYCHAETE FAUNA OF THE GOLD COAST
Fig. 24. Lumbrineris albifrons. Maxillae I to IV from the ventral surface.
Genus HALLA Costa, 1844
Prostomium oval with four eyes and three antennae on its posterior border.
There are two anterior apodous and achaetous segments and no tentacular cirri.
The dorsal cirri are large and foliaceous ; all chaetae are simple and acicular chaetae
are present. Maxillae asymmetrical with two very long maxillary supports, (Text-
fig. 250
Halla parthenopeia (Delle Chiaje), 1828
Holla parthenopeia, Fauvel, 1923, pp. 426-427, fig. 169, a-h, from Naples, Genes and Cadiz.
Locality. Off Accra, Buchanan Survey, in 3-6 to ii'o metres depth (3), two
anterior and one posterior pieces, between 85 and no mm. long.
Collector's note : " This worm is brick-red in life, blue in preservative. Grows
up to 25 in. (625 mm.) long."
The specimens agree very well with Fauvel's description. The superior dorsal
chaetae, however, are not as hispid as presented in his figure. Maxillary plates are
illustrated in Text-fig. 25 and show the plates to have the following teeth : I 11-7,
II 12-15, III 10-9, IV ii-ii, V 1-1. There is, however, variation in this character
and in the other specimens here examined the formulae is : I n-8, II 12-13, III
16-12 +, IV 17-12, V 1-1. Many of the teeth are frequently broken, making accurate
counting difficult. A dorsal incision was made in the pharynx in order to prepare
Text-fig. 25, and the sets of teeth turned outwards. It appears, therefore, as if
maxillae I and II of the right hand side worked against each other. In effect this
maxillae I, with the basal portion of II, bites against I of the left hand side during
mastication.
The large, foliaceous, dorsal cirri each bear an external notch basally, and the
chaetae project laterally between the parapodial lobe and a secondary, anterior
pedal flap.
THE POLYCHAETE FAUNA OF THE GOLD COAST 121
Distribution. This appears to be the first record of this species in the Atlantic
Ocean, south of Cadiz.
Family Orbiniidae
Body divided into two regions : (a) thorax, short and flat with few segments,
neuropodia flattened pads with or without stout bristles ; (b) abdomen, long and
cylindrical, with numerous segments, neuropodia erect. Prostomium without ap-
^=^j c§7^
Fig. 25. H alia par thenopeia. Maxillae from the dorsal surface. (See text,
p. 120, for explanation.)
122 THE POLYCHAETE FAUNA OF THE GOLD COAST
pendages. Proboscis unarmed. Parapodia biramous with simple chaetae. Branchiae
dorsal. Transverse rows of papillae often present on the ventral side of a number of
thoracic segments.
Key to Genera
i. Thoracic neuropodia with vertical rows of pedal papillae .... Orbinia.
Thoracic neuropodia without vertical rows of papillae or with only 2-3 . . . 2.
2. Thoracic neuropodial chaetae entirely slender, pointed .... Haploscoloplos. 1
Thoracic neuropodial chaetae include pointed and hook-like ones . . . Scoloplos.
1 Not recorded from the Gold Coast.
Genus ORBINIA Quatrefages, 1865
Prostomium conical. One pair of lanceolate gills on each segment except a few
anterior ones. Thoracic notopodia with serrated capillary chaetae; neuropodia pad-
like with stout chaetae and papillae. Abdominal notopodia with capillary and forked
chaetae ; neuropodia bilobed with capillary chaetae.
Orbinia foetida var. linguistica (Orlandi), 1896
A ricia foetida var. linguistica, Fauvel, 1927a, pp. 14-16, fig. 4, a-l.
Locality. Off Accra, Buchanan Survey, in 3-6 to n-o metres (1), incomplete,
26 mm. long.
There are 22 segments in the thorax, branchiae begin on the 7th and ventral
papillae appear on the 16th segment, with one pair and gradually increase to five
pairs on segment 21 ; they are absent from segment 22. There are eight papillae
on the posterior neuropodial lobe. Large lanceolate chaetae appear on the nth
to the last thoracic chaetiger. Between rami on abdominal segments there are pro-
nounced cirri.
Distribution. This variety appears to have been recorded only from the Mediter-
ranean Sea, but the stem species is widely known from the English Channel and
Atlantic Ocean and has been collected at Madagascar.
Genus SCOLOPLOS Blainville, 1820
Similar to Orbinia, but without pedal papillae or with only very few, 1-3, and
never with inter-ramal cirri.
Key to Species
1. With capillary chaetae in anterior thoracic neuropodia . . . . . . 2.
Without capillary chaetae in anterior thoracic neuropodia ; with 23 thoracic seg-
ments, with branchiae beginning at the 6th ; with 3 rows of hooks in thoracic
neuropodia ........... 5. johnstonei. 1
2. With pedal papillae on anterior thoracic neuropodia ; with 12 to 20 thoracic seg-
ments, with branchiae beginning at the 9th to 17th. ... 5. armiger. 1
Without pedal papillae on anterior thoracic neuropodia ..... J.
3. Branchiae begin on anterior thoracic segments ..... 5. dubia sp. n.
Branchiae begin on posterior thoracic or anterior abdominal segments
5. madagascarensis.
1 Not recorded from the Gold Coast.
THE POLYCHAETE FAUNA OF THE GOLD COAST 123
Scoloplos madagascarensis Fauvel, 1919
Scoloplos madagascarensis Fauvel, 1919, pp. 433-434, pi. 17, figs. 81-86.
Locality. Off Accra, Buchanan Survey, in 3-6 to n-o metres (2), anterior pieces,
each 33 mm. long ; there are also two median pieces which may be detached portions
of the anterior parts.
In one of the anterior pieces the thorax is made up of 21 segments, of which the
first nine have two rows of neuropodial hooks and the remainder one. Branchiae
appear on the last thoracic segment, on which there is also one pedal papilla. In the
other anterior piece there are 26 thoracic segments, all of which have one row of
neuropodial hooks. Branchiae appear on the first abdominal segment. This piece
is without pedal papillae.
Both specimens have a few short forked chaetae in abdominal notopodia, one arm
of the fork being about twice as long as the other. These chaetae are very easily
broken and difficult to find.
Fauvel (1919) originally described 5. madagascarensis with a thorax of 26-27
segments and with branchiae appearing at the 21st or 22nd segments becoming
fully developed at the 25th or 26th. It will be seen that the specimens from Accra
agree in one or other of these characters, and it is suggested that the species may be
more variable than has hitherto been indicated.
Distribution. Previously recorded from Madagascar and Inhaca Island, Delgoa
Bay, Transvaal (Day, 1951).
Scoloplos dubia sp. n.
Holotype ; B.M. (N.H.) Reg. No. 1953.3. 1 .1230.
Locality. Off Accra, Buchanan Survey, in 3-6 to ii-o metres depth, complete,
about 90 mm. long.
The sharply pointed prostomium is without visible eyes. There are 22 thoracic
segments. Thoracic neuropodial capillaries (Text-fig. 26, a) appear on every foot ;
notopodial chaetae throughout the body are similar to these. Forked notopodial
chaetae have not been seen. Thoracic neuropodial hooks are shown in Text-fig.
26, b, they occur in three to four rows on each segment. In the posterior region of
the abdomen the neuropodial aciculae become external and hooked (Text-fig. 26, c).
Paratypes ; B.M. (N.H.) Reg. No. 1953.3. 1 . 1231-4, from the same locality as the holotype.
These specimens are all incomplete, the longest measuring 65 mm. In all of them
the branchiae appear at the 7th foot ; in two the thorax has 22 segments, in one
21 and in the other 23. In other characters they are identical with the holotype.
As suggested by its name the status of this new species is doubtful. It is un-
doubtedly closely related to S. johnstonei, from which it is distinguished principally
in having thoracic neuropodial capillaries and posterior hooked aciculae. Examina-
tion of material from areas between the Gold Coast and the Cape of Good Hope (the
distributional zone of 5. johnstonei) may clarify the relationship.
ZOOL. 3, 2. IO
124
THE POLYCHAETE FAUNA OF THE GOLD COAST
Fig. 26.
0-5mrn
Scoloplos dubia sp. n. (a) Thoracic neuropodial chaeta, (b) thoracic neuropodial
hook, (c) posterior abdominal parapodium.
Family Spionidae
Body not divided into distinct regions. Prostomium without antennae ; (some-
times with frontal peaks) ; with palps and eyes. Biramous parapodia with foliaceous
ventral and dorsal cirri ; dorsal branchiae on a number of segments. Chaetae simple
capillaries and hooded hooks.
Genus PRIONOSPIO Malmgren, 1867
Prostomium without frontal peaks. Branchiae 3-1 1 pairs, often pinnate, confined
to anterior segments. Simple capillaries and pluridentate hooded hooks are present.
Prionospio pinnata Ehlers, 1901
Prionospio africana, Augener, 1918, pp. 402-5, pi. 6, figs. 162-3, text-fig. 51.
Prionospio pinnata, Fauvel, 1932, p. 173.
Localities. Off Accra, Stn. 69 (2), incomplete, both 7 mm. in length ; Buchanan
Survey, off Accra, 3-6 to n-o metres, depth (7), one complete, 55 mm. long, others
incomplete 10-41 mm. long.
Pinnate branchiae, if present, occur on the first three chaetigers only, frequently
they have been accidentally lost. Several of the specimens from the Buchanan
THE POLYCHAETE FAUNA OF THE GOLD COAST 125
Survey still retain one of the long pair of palps, which extends from the posterior
region of the prostomium to the 16th chaetiger. The palps are somewhat curved
and grooved dorsally (Text-fig. 27). Their external margin is marked with mauve
Fig. 27. Prionospio pinnata. Grooved palp from the left hand side of the prostomium.
(Specimen from the Buchanan Survey.)
vertical stripes. Four eyes are set in a trapezium on the prostomium, which is
enclosed between two membranous wings. Within these wings the prostomium
consists of a prominent longitudinal ridge.
Distribution. Atlantic Ocean, Indian Ocean and Pacific Ocean.
Family Cirratulidae
Body short, cylindrical, with numerous segments ; head without appendages, with
or without eyes. Proboscis smooth, unarmed. Dorsal palps or tentacular cirri on
an anterior segment ; long slender branchiae inserted above the notopodia ; para-
podia biramous ; chaetae simple capillaries and acicular hooks.
Key to Genera
1. Branchiae appearing on the same segment as the tentacular cirri . . . Cirratnlus.
2. Branchiae appearing on a few segments in front of the one bearing tentacular cirri
Audoninia.
Genus AUDOUINIA Quatrefages, 1865
First three segments achaetous. Branchiae occur on the first chaetigers to nearly
the last ones. Of the three species of this genus recorded here, each is confined to a
particular area of the Gold Coast. Thus A. punctata has been found only in the
Tenpobo area ; A. tentaculata only on the Winneba shore, and A. filigera only in
the Sekondi-Axim region. No overlap has been noted.
Key to Species
1 . Branchiae in the middle body region situated half the distance above the notopodium
that the notopodium is above the neuropodium . . . A. tentaculata.
Branchiae in the middle body region situated much more than half the distance
above the notopodium than the notopodium is above the neuropodium . . 2.
126 THE POLYCHAETE FAUNA OF THE GOLD COAST
2. One hook in the neuropodium in the posterior part of the body. Branchiae in the
middle body region situated, normally, much further than twice as far above the
notopodium as the notopodium is above the neuropodium . . . . A . filigeva.
Three or more hooks in the neuropodium in the posterior part of the body.
Branchiae in the middle body region situated at least as far above the
notopodium as the notopodium is above the neuropodium . . . A . punctata.
Audouinia tentaculata (Montagu), 1808
Cirratulus tentaculatus var. meridionalis, Augener, 1918, pp. 461-463, pi. 6, fig. 175 — 176.
Audouinia tentaculata, Fauvel, 1927a, pp. 91-92, fig. 32, a-g.
Locality. Winneba shore, 22.xi.49 (20), between 18-35 mm. long, and another
ten specimens, badly mutilated, in association with tubes of Sabellaria eupomatoides
from this area collected on the same day.
The tentacular cirri appear between the 6th and 7th chaetigers. Typically there
are two hooks in the neuropodia over the greater part of the body, though for some
segments after the 12th foot there may be three to four.
Distribution. Common in the European zone from the North Sea, Irish Sea,
English Channel and the Atlantic Ocean. Known also from German South- West
Africa (Augener, 1918) and the Cape (Day, 1951).
Audouinia filiger a (Delle Chiaje), 1841
Audouinia filigeva, Fauvel, 1927a, pp. 92-93, fig. 32, h-m.
Audouinia filigeva, Fauvel, 1932, p. 178.
Localities, (i) Sekondi, high level plateau, 21.ii.49 (4), complete, up to 20 mm.
long. (2) Dixcove, shore (12), mutilated, 20-40 mm. long. (3) Princes Town, shore,
15.iv.49 (4), complete, 20-25 mm. long. (4) Axim, Hospital Reef (3), 10-12 mm.
long.
The tentacular cirri occur between the 4th and 5th chaetigers. After the appearance
of two neuropodial hooks at the 12th foot, which persist for a small number of seg-
ments, there is a reduction to one, which condition is maintained up to the end of
the body.
Distribution. Well known from the Atlantic Ocean, Persian Gulf, and the Indian
and Pacific Oceans.
Audouinia punctata (Grube), 1858
Civvatulus punctatus, Augener, 1918, pp. 465-467.
Audouinia punctata, Day, 1951, p. 47.
Locality. Tenpobo shore, approximately 370 specimens, measuring up to 20 mm.
long were collected here during February, 1950.
The tentacular cirri are situated between the 3rd and 4th chaetigers. There are
normally three hooks in the neuropodium over the greater part of the body, but
anteriorly (i.e. just after the 12th parapodia) there may be a reduction to two and
posteriorly an increase to four. Colour in spirit grey-black, flecked with black, but
considerable areas of white may be present ; white to grey tentacular cirri with
THE POLYCHAETE FAUNA OF THE GOLD COAST 127
black lateral stripes ; gills grey black with sometimes an orange tip ; young forms
brown.
Distribution. This species is known from the Atlantic, south of Florida to the
Cape of Good Hope, and from the Natal coast.
Genus CIRRATULUS Lamarck, 1818
Similar to Audouinia except as in above Key.
Cirratulus filiformis Keferstein, 1862
Cirratulus filiformis Fauvel 1927a, pp. 94-95, fig. 33, h.
Locality. Off Accra, Buchanan Survey, in 3-6 to n-o metres (5), all incomplete,
the largest piece measuring 38-0 mm. long.
The prostomium is sharply pointed, and has no eyes. On the 1st chaetiger there
is one pair of long and filiform branchiae and these are present up to the end of
the body. All chaetae are simple capillaries, there are no hooks present. Anteriorly
segments are narrow ; posteriorly they are long.
Distribution. This species is well known from West European waters and has
been reported from Senegal by Fauvel (1902).
Family Flabelligeridae
Prostomium and peristomium with eyes, palps and branchiae, completely re-
tracted into the oral aperture, made visible only by dissection. Chaetae of anterior
segments long, directed forwards to form a cephalic cage. Parapodia biramous and
normally without distinct processes. Notopodial chaetae simple ; neuropodial
chaetae sigmoid or hooked or compound with a sickle-shaped terminal piece.
Genus STYLARIOIDES Delle Chiaje, 1828
Body elongated, with numerous papillae on its surface. Prostomium with two
stout palps. Branchiae filiform, on a peduncle, retractile. Notopodial chaetae
simple capillaries, annulated ; neuropodia with simple capillaries and sometimes
with strong hooks.
Key to Species
1. Neuropodial chaetae long and slender throughout . . . .5. scutigeroides.
Some neuropodial chaetae hooks . . . . . . . . . 2.
2. Neuropodial hooks with bidentate tips ...... 5. arenosus.
Neuropodial hooks with entire tips ....... 5. tropicus.
Stylarioides scutigeroides Augener, 1918
Stylarioides scutigeroides Augener, 191 8, pp. 444-447, pi. 6, figs. 155 and 185, text-fig. 66.
Localities. Off Accra : Stn. 55 (1) ; Stn. 69 (4) ; Buchanan Survey, in 3*6 to
ii- o metres (5), only four of these specimens are complete, measuring between
15 and 30 mm. in length.
128 THE POLYCHAETE FAUNA OF THE GOLD COAST
The complete body consists of 70 parapodia. The 1st chaetiger is telescoped into
the 2nd, the 2nd into the 3rd and the 3rd into the 4th. Thereafter the segments are
separated only by intersegmental grooves anteriorly and ridges posteriorly.
Chaetae of the first parapodia vary between 2-5 to 4-5 mm. in length (Text-fig. 28,
a and b) and are directed forward, forming the characteristic cephalic cage of the
Flabelligeridae. Chaetae of the second parapodia are between 1 and 2-5 mm. long
and are also directed anteriorly. Thereafter the chaetae decrease in size and project
laterally. Notopodia and neuropodia in the first and second parapodia are close
together, elsewhere they are far apart. The anterior region of the body is swollen,
posteriorly it narrows, so that in one specimen the width across the 12th segment is
2-5 mm. and across the 30th, 0-5 mm. Except for a few posterior segments the body
is heavily encrusted with sand grains.
All chaetae are long and slender ; notopodial bristles are illustrated in Text-fig.
28, d, neuropodial bristles in Text-fig. 28, c. Papillae are present all over the body
but are hidden to superficial view by the heavy encrustation of sand. On the first
and second parapodia a single finger-shaped papilla, which may be bi- or trifid
terminally, is associated with each group of chaetae, and there are sometimes one
or two interramal in position. On the 3rd and 4th chaetigers simple finger-shaped
papillae are present between the rami, encircling the segments like coronets. There
are 6-7 dorsally between the notopodia, 2-3 laterally between the notopodia and
neuropodia and 6-7 ventrally between the neuropodia. After the 5th segment the
papillae become smaller and more scattered in arrangement. Posteriorly they are
less numerous.
Distribution. 5. scutigeroides is known only from the Gold Coast, the French
Congo and Cabinda.
Stylarioides arenosus (Kinberg), 1867
Pycnoderma fernandense, Augener, 1918, pp. 448-452, pi. 6, figs. 148, 182 ; pi. 7, figs. 237,
238 ; text-fig. 67, from Saltpond, Gold Coast.
This species has not been collected in the present survey. Stout neuropodial
chaetae, with bidentate tips appear at the 5th chaetiger. The secondary tooth at
the tip is easily broken and the chaetae have frequently been described as unidentate.
Papillae run along the body in longitudinal rows between the rami. Normally there
are between 10 to 12 to each segment, four ventral and four dorsal and one or two
laterally. One of Augener's largest specimens measured 39 mm. long for 75 segments,
but Hartman (1948) found one of Kinberg's types measured 60 mm. for 70 segments.
Distribution. Around the coast of Africa from the Gold Coast to Natal.
Stylarioides tropicus Augener, 191 8
Stylarioides tropicus Augener, 1918, pp. 437-440, pi. 7, figs. 220, 221, text-fig. 63.
This species has not been collected in the present survey. It appears to be charac-
terized by strong hooks with entire tips which appear at the 4th chaetiger, otherwise
Augener's*" description is difficult to follow. One of Augener's specimens measured
24 mm. long fori 32 segments.
THE POLYCHAETE FAUNA OF THE GOLD COAST
129
<<■>
Fig. 28. Stylariodes scutigeroides. Chaetae ; (a) and (b) of the cephalic cage
(c) neuropodial ; (d) notopodial.
130 THE POLYCHAETE FAUNA OF THE GOLD COAST
Distribution. The species was collected from Liberia, the Gold Coast, Accra,
French Congo, Cabinda and Angola.
Family Opheliidae
Body short, convex dorsally, with or without a gutter ventrally. Prostomium
without appendages, frequently with cephalic eyes. Sometimes with lateral eye
spots. Biramous parapodia with simple chaetae. Pygidium with papillae or some-
times prolonged into an anal tube.
Key to Genera
i. With lateral branchiae .......... Armandia.
2. Without lateral branchiae ......... Polyophthalmus .
Genus ARMANDIA Filippi, 1861
Body not divided into distinct regions ; a deep median and two lateral ventral
grooves. Branchiae from the 2nd chaetiger to the end of the body. Pygidium with
anal funnel fringed with papillae and a median cirrus. Lateral eye-spots.
Armandia intermedia Fauvel, 1902
Armandia intermedia Fauvel, 1902, pp. 86-89, figs. 29, 30, from Senegal.
Armandia intermedia, Augener, 1918, pp. 424-425, from Senegal and Angola.
Armandia intermedia, Day, 1949, p. 449, from St. Helena.
Locality. Off Accra, Stn. 69 (1), 12 mm. long.
There are 29 chaetigers, the 1st abranchiate, the next 25 branchiate (though some
of the posterior branchiae have fallen off, the point of attachment to the superior
edge of the parapodia can be seen), and the last three abranchiate. Thirteen pairs
of lateral eye-spots are situated on the 7th to the 19th chaetigers. There are three
cephalic, subcutaneous, eye-spots disposed in a triangle. The anal funnel is fringed
with numerous papillae, but many have fallen off and the exact number cannot be
estimated.
Distribution. This species is only known through the above records.
Genus POLYOPHTHALMUS Quatrefages, 1850
Similar to Armandia but with no branchiae.
Polyophthalmus pictus (Dujardin) 1839
Polyophthalmus pictus, Fauvel, 1927a, pp. 137-138, fig. 48, l-o.
Localities, (i) Tenpobo shore, 6.ii.5o (2), 6-8 mm. long ; (2), Lighthouse Reef,
Axim, 8.L51 (1), 9 mm. long.
There are 28 chaetigers with about 11 pairs of eye spots on the 6th to 16th seg-
ments, but these are very difficult to see and may vary considerably in position.
THE POLYCHAETE FAUNA OF THE GOLD COAST 131
Distribution. English Channel, Atlantic, Mediterranean, and the Red Sea ;
Pacific and Indian Oceans.
Family Maldanidae
Body of few segments some of which are exceptionally long. Prostomium without
appendages, often with a rimmed cephalic plate and a median keel, on each side
of which is a nuchal groove. Buccal segment achaetous. Parapodia biramous,
notopodial chaetae capillaries, neuropodial chaetae uncinate ; no parapodial cirri.
Anal segment with funnel and cirri or a flat plate. Inhabitants of cylindrical tubes
which may be thin and covered with sand or thick and coated with mud.
Genus MALDANE Grube, i860
Cephalic keel arched, rim divided into three by two lateral notches. Anus dorsal
with a slanting anal plate. Ante-anal segments achaetous. Neuropodial chaetae
missing from the first segment. Notopodial chaetae of three kinds. Tube coated
with mud.
Maldane sarsi Malmgren, 1865
Maldane sarsi, Fauvel, 1927a, pp. 197-199, fig. 69.
Locality. Off Accra, Buchanan Survey, in 36-5 metres off Castle (1), complete,
about 62 mm. long.
Prostomium with the cephalic keel strongly arched and the rim smooth, but
notched on each side.
The nuchal grooves are short, curved and deep. There are 19 chaetigers and two
achaetous ante-anal segments. Notopodial chaetae are of three types : (a) long
capillaries, barbed along their entire length ; (b) limbate capillaries, slightly barbed
at their tips ; (c) very short geniculate capillaries with a limb on their convex borders.
Uncini with one very large tooth surmounted by 5-6 rows of smaller teeth ;
beneath the large tooth there is a cluster of fine hairs.
Distribution. Cosmopolitan.
Family Oweniidae
Cylindrical body of few segments having the prostomium fused with the buccal
segment and being either devoid of appendages or terminating in a membranous
lobe. Notopodial chaetae simple capillaries ; neuropodial chaetae uncinate, very
small, with a bent hooked tip. Tube coated with sand and shell fragments.
Genus OWENIA Delle Chiaje, 1841
Prostomium terminating in a membranous branchial lobe. Buccal segment
achaetous ; the first three chaetigers without uncini. Notopodial chaetae slender,
slightly spinous. Pygidium bilobed.
132 THE POLYCHAETE FAUNA OF THE GOLD COAST
Owenia fusiformis Delle Chiaje, 1841
Owenia fusiformis, Fauvel, 1927a, pp. 203-204, fig. 71, a-f.
Owenia fusiformis, Augener, 19x8, pp. 492-493.
Localities. Off Accra : (a) Stn. 5 (25) ; Stn. 11 (4) ; Stn. 32 (numerous) ; Stn.
53 (24) ; Stn. 54 (numerous) ; Stn. 55 (50 +) ; Stn. 61 (7) ; Stn. 65 (numerous) ;
Stn. 66 (numerous) ; Stn. 67 (36) ; Stn. 121 (2) ; Buchanan Survey, 3-6 to n-o
metres (9) ; (b) Dredge haul off Chorkor, 1 .xi.50 (20).
The animals vary in length between 40-60 mm., the tubes, many of which are
empty, particularly from Stns. 55, 65 and 66, measure up to 88 mm. The thorax
consists of the achaetous buccal segment and the first three chaetigers which have
only notopodial capillaries ; there is no neuropodial component in this region. The
remaining segments, which are all biramous, make up the abdomen, and are long
in the anterior region, but posteriorly become progressively smaller. The tubes
consist of agglutinated small shells and sand, the former orientated at right angles
to the long axis, and are grey in colour.
Distribution. Cosmopolitan.
Family Sabellariidae
Body divided into four regions, (a) an anterior region with opercular disc and
stalk, (b) a thorax of two anterior and three or four parathoracic segments, (c) an
abdomen with uncinigerous dorsal rami and ventral rami with capillaries, (d) a
caudal region, unsegmented, achaetous and apodous. The prostomium is hidden
between the two large opercular stalks which carry paleae in concentric rows. Two
palps. Simple branchiae are carried on the thorax and abdomen. Tubes are laid
down and normally form sandy reefs of varying extent ; occasionally species may
be solitary.
Genus SA BELLA RIA Lamarck, 1818
The operculum is short, with numerous filiform tentacles on the ventral side and
provided with three visible rows of paleae. (The middle and inner rows probably
represent a single row.) Thorax with two anterior segments, having only capillary
chaetae, and three parathoracic segments with flattened paleae dorsally and similar,
smaller chaetae, ventrally.
Key to Species
1. Animal small, less than 10 mm. in length when sexually mature ; tubes made up of
small particles of sand . . . . . . . . .5. eupomatoides.
Animals large, more than 15 mm. in length when sexually mature ; tubes made up
of large and coarse sand grains . . . . . . . . . 2.
2. Central tooth of outer paleae bluntly spatulate, entire or bifid 5. spinulosa var. intoshi.
Central tooth of outer paleae with a barbed point , , 5. spinulosa var. alcocki.
THE POLYCHAETE FAUNA OF THE GOLD COAST
133
Sabellaria eupomatoides Augener, 1918
Sabellaria eupomatoides Augener, 1918, pp. 503-504, pi. 7, figs. 187-190, text-fig. 83, from
Angola.
Localities. Numerous masses of the tubes of this species were collected from
Winneba shore, Apam shore and Christiansborg shore at various times in 1949 ;
specimens normally measure between 5-8 mm. in length.
This species is characterized by its small size (a mature female from Winneba,
collected 22.xi.49, measured 7 mm. long) and the form of the opercular paleae
(Text-fig. 29). In some specimens several paleae of the median row project above
Fig. 29. Sabellaria eupomatoides.
(b) enlarged terminal piece of (a)
row. (Specimen from Winneba.)
Opercular paleae : (a) from the external row ;
(c) from the outer row and (d) from the median
the level of the others. There are nine pairs of simple gills from the 2nd thoracic
segment backwards.
S. eupomantoides was described by Augener in 1918 from specimens collected
without their tubes, and has not been recorded since then except as above.
Sabellaria spinulosa Leuckart, 1849
Acicular chaetae are present on the dorsal face of the operculum, immediately
posterior to the terminal crown of paleae. There is an inter-peduncular cirrus.
Paleae in the external row have a barbed median tooth.
134 THE POLYCHAETE FAUNA OF THE GOLD COAST
Sabellaria spinulosa var intoshi Fauvel, 1914
Sabellaria spinulosa var. intoshi Fauvel, 1927a, pp. 208-211, fig. 73^.
Localities, (i) Tenpobo shore, about 20 specimens and tubes were collected in
January and February, 1949, several were complete and measured up to 36 mm.
long without the tail. (2) Apam (1), incomplete, 20 mm. long, and numerous tubes.
(3) Dixcove shore (1), incomplete, 30 mm. long, with tube. (4) Axim Lighthouse
Reef, 14.iv.49 (4), up to 38 mm. long, with tubes.
This variety differs from the stem species in having the median tooth of paleae
of the outer row with a spatulate tip. Occasionally this tip may be bifid.
Distribution. English Channel and the Atlantic ocean (Gulf of Guinea).
Sabellaria spinulosa var alcocki Gravier, 1906
Sabellaria spinulosa var alcocki, Fauvel, 1927a, pp. 208-211, fig. 73, k-m.
Locality : Axim, Hospital Reef, 7.L51 (1), 18 mm. long, with tube.
Some of the paleae in the median row are always erect in this variety, otherwise
it is similar to the stem species.
Distribution. English Channel, North Sea, Atlantic Ocean, Indian Ocean and
the Red Sea.
Family Sternaspidae
The form adopted in the family is aberrant. The body is swollen at both ends ;
the prostomium small and devoid of appendages ; the first three segments have
chaetae, the middle ones are achaetous, and posteriorly there is a ventral shield
from the borders of which chaetae radiate. Branchiae are posterior to the shield.
Genus STERN ASPIS Otto, 1820
Branchiae filiform, set in two bundles.
Sternaspis scutata (Ranzani) var. africana Augener, 1918
Sternaspis fossor Stimpson var. africana Augener, 1918, pp. 608-613, text-fig. 109.
Sternaspis scutata var. africana, Monro, 1930, pp. 179-180.
Localities. Off Accra : Stn. 27 (42) ; Stn. 28 (2) ; Stn. 33 (6) ; Stn. 47 (2) ;
Stn. 59 (1) ; Stn. 61 (8) ; Stn. 68 (14) ; Stn. 69 (19) ; Stn. 71 (3) ; Stn. 130 (1) ;
Buchanan Survey, 3*6 to iro. metres (4).
The specimens vary between 5 and 15 mm. in length, but these measurements must
be treated with caution, for in many cases the anterior chaetigers have contracted
into the succeeding segments.
As Augener (1918) and Monro (1930) pointed out, the variety is distinguished,
from the stem species by the greater number of chaetae (about 20 as opposed to
10-12) which occur on the anterior segments, The chaetae are also more slender in
THE POLYCHAETE FAUNA OF THE GOLD COAST 135
the variety. These small differences seem to divide clearly the West African forms
from the European and Antarctic examples, though further examination may
indicate that the difference is merely ecological. Thus S. scutata has been reported
from 68-1080 metres (Monro, 1930), from about 300 metres (Chamberlin, 1919),
and S.fossor from 68-130 metres, (Fauvel, 1936) and from 120-130 metres, (Takahasi,
1938), whereas S. scutata var. africana has not been reported beyond a depth of
67 metres. Unfortunately many scattered records of the parent species do not give
depth details or particulars of the morphological differences noted above, so that
an accurate consideration of this issue is not possible. However, further collecting,
at depth, off the Gold Coast may clarify the matter, and if it is confirmed that the
morphological differences are non-genetic, but induced by the ecological conditions,
the taxonomic treatment must be modified. If they prove to be genetic and asso-
ciated with geography africana must rank as a subspecies.
Distribution. 5. scutata var. africana is only known from off the W. African
coast, from Senegal south to Angola.
Family Ampharetidae
Body divided into two regions, (a) thorax with notopodial capillaries and neuro-
podial uncini, (b) abdomen with neuropodial uncini only. Tentacles retractile into
the mouth. There are three or four pairs of branchiae on anterior segments and they
may be subulate, pectinate or bi-pectinate. With or without groups of paleae.
Key to Genera
i. With paleae and without dorsal hooks behind the branchiae . . . . . 2.
Without paleae and with dorsal hooks behind the branchiae .... Isolda.
2. With one of the four pairs of branchiae pectinate ; 17 thoracic chaetigers Phyllamphicteis.
With one of the four pairs of branchiae bi-pectinate ; 16 thoracic chaetigers Pterolysippe.
Genus PHYLLAMPHICTEIS Augener, 1918
With two fan-like groups of paleae ; thorax of 17 chaetigers with uncini appearing
at the 4th. Abdomen of 15 chaetigers. Four pairs of branchiae, three subulate, one
pectinate.
Phyllamphicteis collar ibranchis Augener, 1918
Phyllamphicteis collaribranchis Augener, 1918, pp. 509-512, pi. 7, figs. 239-241, text-fig. 85,
from Wappu on the Ivory Coast.
Localities. Off Accra : Stn. 28 (1) ; Stn. 33 (1) ; Stn. 69 (1) ; Stn. 70 (1) ;
Stn. 97 (4) ; Stn. in (1) ; several of these specimens are complete measuring up to
20 mm. long.
There are 8-10 paleae in each fan. Thoracic uncini have five teeth ; notopodial
capillaries are slightly limbate. Both parapodial rami have small cirri on thorax
and abdomen. The tubes are of fine dark grey mud with a few adhering shell particles.
Four specimens collected in the Buchanan Survey, in 3-6 to iro metres off Accra
136 THE POLYCHAETE FAUNA OF THE GOLD COAST
and measuring between 20 and 30 mm. long, differ from the above in having two
pairs of pectinate branchiae instead of one, though similar in other respects. It
may be that they represent a different species, but it is possible that there is a
considerable range of variation in this character within the species. Additional
material and observations are needed to settle this point.
Distribution. Only known from the above records.
Genus PTEROLYSIPPE Augener, 1918
With two groups of paleae, each like a fan. Thorax of 16 chaetigers with uncini
appearing at the 4th. Four pairs of branchiae, 3 subulate and the other bi-pectinate.
Pterolysippe bipennata Augener, 1918
Pterolysippe bipennata Augener, 1918, pp. 512-514, plate 6, figs. 173 & 174, text fig. 86, from
Whydah, Dahomey.
Localities. Off Accra: Stn. 59 (1), with tubes; Stn. 133 (1) ; Buchanan Survey,
in 3-6 to ii- metres (4), and in 41-86 metres off the Lagoon (1) ; several of these
specimens are complete and measure up to 18 mm. long.
The prominent bi-pectinate gill is coloured with numerous small purple marks.
There are 15 abdominal segments. Thoracic uncini have 5-6 teeth, notopodial
chaetae are limbate. Branchiae become easily detached in this species and it is
rare to find a specimen with the complete set. No cirri are visible on the parapodia.
Tubes are thick, of fine grey-black mud.
Distribution. Only known through the above records.
Genus ISOLD A Muller, 1858
Without paleae. Number of thoracic and abdominal segments variable. Four
pairs of branchiae, two subulate, and two feather-like. Two dorsal anterior hooks,
immediately behind the branchiae.
Isolda whydahensis Augener, 1918.
Isolda whydahensis Augener, 1918, pp. 514-518, pi. 7, fig. 216, text-fig. 87.
Locality. Off Accra, Buchanan Survey, in 5-5 metres off the breakwater (i),
incomplete, length 15 mm.
There are 16 thoracic chaetigers the first three of which are very difficult to see.
They lie on the prostomial collar and in the specimen examined the chaetae had
been broken off. Neuropodial uncini appear at the 5th chaetiger, are short and have
five teeth. Thoracic capillaries are limbate. The transverse membrane which, with
the prostomial collar, encloses the branchiae and dorsal hooks, is smooth. Both of
the inner pairs of branchiae are feather-like, the outer pairs are subulate.
Distribution. I. whydahensis is known also from Whydah, Dahomey, the French
Congo and Angola.
THE POLYCHAETE FAUNA OF THE GOLD COAST 137
Family Terebellidae
Body with thorax having notopodial capillary chaetae on all chaetigers and
neuropodial uncini on some, and abdomen with neuropodial uncinionly. Prostomium
with non-retractile tentacles. Branchiae, when present, normally on the anterior
segments. In the thorax there are ventral glandular scutes. The tubes are coated
with sand.
Key to Genera
1. Without branchiae ........... Amaea
With branchiae ............ 2
2. With a single quadripartite pectinate gill ...... Terebellides
With more than one gill ........... 3
3. Thorax with more than 17 chaetigers ....... Streblosoma
Thorax with 17 chaetigers or less ......... 4
4. Uncini pectinate ........... Loimia
Uncini avicular ............ Pista
Genus PISTA Malmgren 1865
Thorax with 15-17 chaetigers ; eye-spots may be present. One, two or three pairs
of branchiae. Notopodial chaetae smooth ; neuropodial uncini appear in the thorax
on the 2nd chaetiger.
Pista grubei Augener, 1918
Pista grubei, Augener 1918, pp. 542-5, pi. 7, fig. 252-3, text-fig. 92.
Localities. Off Accra: Stn. 28 (1) ; Stn. 61 (1) ; Stn. 69 (1) ; Stn. 131 (1).
None of the specimens is complete, they vary between 25 and 47 mm. in length.
There are 17 thoracic chaetigers, of which the first has only simple notopodial
chaetae. In the next six segments there are also neuropodial groups of uncini in
single rows, all the hooks lying in the same direction. The remaining 10 thoracic
chaetigers, however, have the uncini lying alternately head to tail in a single row.
The uncini are avicular in shape and all have a projecting basal process.
The gills are in two pairs, consisting of a long basal trunk supporting a mass of
whorled branching filaments. They are dorsal, anterior to the first chaetiger.
Distribution. Known also from Senegal, Ivory Coast, and French Congo.
Genus TEREBELLIDES Sars, 1835
With a single gill divided into four pectinate arms. Notopodial chaetae long,
tapering and winged. Uncini of two types, the thoracic elongated, acicular, the
abdominal pectinate.
Terebellides stroemi Sars, 1835
Terebellides stroemi, Fauvel, 1927a, pp. 291-2, fig. 100, i-g.
Terebellides stroemi, Fauvel, 1932, pp. 234-5.
Localities. Off Accra : Stn. 28 (4) ; Stn. 33 (2) ; Stn. 59 (4) ; Stn. 73 (2) ;
Stn. 133 (2). Most of these specimens are complete, measuring between 10 and
30 mm. long. Buchanan Survey, 36-5 m. off Castle (1), 35 mm. in length.
138 THE POLYCHAETE FAUNA OF THE GOLD COAST
Body rarely with more than 60 segments, the first 18 being thoracic, with the gill
as above. Segments 3 to 6 have their anterior border free. Notopodial chaetae are
simple and begin on segment 3 ; neuropodial uncini appear at the 6th chaetiger and
are long, geniculate, acidular hooks ; the remaining thoracic segments also have
long uncini, but these end in blunt tips surmounted by small denticles. The
abdominal uncini are avicular with a short base and transverse rows of teeth above
the main fang.
Distribution. Mediterranean Sea, Atlantic Ocean, Indian and Pacific Oceans.
Genus STREBLOSOMA Sars, 1872
Body with a large number of segments ; often with eyes ; with two or three pairs
of branchiae (sometimes five?) on the 1st, 2nd, or 3rd chaetigers. Notopodial
capillary bristles appear on the 2nd segment (1st branchiate). Neuropodial uncini
appear on the 5th segment (4th chaetiger).
Streblosoma persica (Fauvel), 1908
Gvymaea persica, Fauvel, 1911, pp. 419-421, pi. 20, figs. 35-43.
Pseudothelepus nyanganus, Augener, 1918, pp. 552-555, pi. 7, figs. 257, 258, text-fig. 96.
Streblosoma persica, Fauvel, 1950, p. 384.
Localities, (i) Tenpobo shore, about 100 specimens collected in January, 1949,
and February, 195 1, measuring between 20-50 mm. in length. (2) Apam shore,
16.ii.49 (12). (3) Dixcove shore, 7 . ii . 5 1 (12). (4) Axim, 13/14.^.49 (2).
The species is characterized by (a) a semi-circle of eye-spots on the dorsal surface
of the tentacular disc and (b) the shape of the uncini, which have a large basal tooth,
paired smaller median teeth and above these a single smaller tooth with several tiny
ones on either side of it.
Notopodial chaetae occur on almost all segments. There are three pairs of bran-
chiae, formed of numerous filaments, one pair on each side of the first three chaetigers.
The first pair are based on transverse pads which form semi-circles around the anterior
aspects of the notopodia of the first chaetiger. The remaining two are based on pads
situated wholly on the dorsal side of the notopodia.
Distribution. The species has been recorded from the Persian Gulf, the Gulf of
Manaar, the Island of St. Thomas in the Gulf of Guinea, Angola and Senegal.
Genus LOIMIA Malmgren, 1865
Body with 17 chaetigers and three pairs of aborescent gills. Anterior segments
have lateral lobes. Notopodial chaetae winged. Uncini pectinate, opposed back-to-
back in double rows from the 7th to the 17th thoracic chaetiger.
Loimia medusa (Savigny), 1822
Loimia medusa Augener, 1918, pp. 539-541, text-fig. 91.
Lattice fauvelii Day, 1934, PP- 7 I- 73> fig* J 4» a ~d-
Locality. Off Accra, Buchanan Survey, in 36-5 metres off the Castle (1), an
incomplete specimen of 34 chaetigers measuring 28 mm. in length.
THE POLYCHAETE FAUNA OF THE GOLD COAST 139
The three pairs of gills, of which the first is very much the largest, all lie anterior
to the first chae tiger. The uncini have 6 teeth in both thorax and abdomen.
Distribution. The species has been recorded from the English Channel, the
Atlantic, Indian and Pacific Oceans, the Red Sea and the Persian Gulf.
Genus AMAEA Malmgren, 1865
Prostomium large, with many tentacles. Branchiae and eye-spots absent. Thorax
of few segments, with notopodial chaetae. Abdomen consisting of an anterior
achaetous portion and a posterior region with slender uncini.
Amaea accraensis Augener, 1918
Amaea accraensis Augener, 191 8, pp. 561-562, pi. 7, fig. 246, text-fig. 98, from Accra.
This species has not been found in the present survey. The thorax has 11 chae tigers
on the left hand side and 13 on the right. Notopodial chaetae are barbed capillaries.
The achaetous abdominal region is about two-thirds as long as the thorax. There
are 26-27 chaetigers posteriorly on the abdomen, bearing hooks with slightly bent
tips. Augener's specimen measured 18 mm.
Distribution. This species is only known from the type specimen.
Family Sabellidae
Body divided into two regions : (a) thorax, with few segments, having notopodial
capillaries and neuropodial uncini, and (b) abdomen, normally with numerous
segments having notopodial uncini and neuropodial capillaries. Ventral glandular
shields divided by a longitudinal groove. First chaetiger with an entire, or notched
collar. Branchiae encircling the mouth. Tubes of mucous, or membranaceous or
horny, material.
Key to Genera
1. Thoracic neuropodia with a single row of avicular uncini, pickaxe-shaped chaetae
absent .............. 2.
Thoracic neuropodia with a row of avicular hooks and a row of pickaxe-shaped
chaetae (fig. 30, a) . . . . . . . . . . Potamilla.
2. Branchiae without dorsal sty lodes ........ Laonome.
Branchiae with dorsal sty lodes ......... Dasychone.
Genus LAONOME Malmgren, 1865
Two symmetrical, semi-circular, branchial lobes not in the form of a spiral.
Branchial filaments without stylodes. No pickaxe-shaped chaetae.
Laonome punctur at a (Augener), 1918
Demonax puncturatus Augener, 1918, pp. 576-580, pi. 6, fig. 170, 171, text-fig. 102.
Euratella puncturata Monro, 1930, p. 203.
Localities, (i) Accra : considerable quantities of tubes were collected from
Christiansborg shore on 15.1.49, 14.ii.49, and 17. hi. 49. (2) Apam, 16.ii.49,
ZOOL. 3, 2. II
i 4 o THE POLYCHAETE FAUNA OF THE GOLD COAST
numerous specimens and tubes. (3) Axim shore, 13/14.^.49, numerous specimens
but no tubes.
The prominent prostomial collar opens dorsally and forms two triangular lappets
ventrally. There are between 10 and 12 branchial filaments which, in most specimens,
bear at least one distinct pigment band. Pigment spots are also present on the
collar and between the rami of thoracic and abdominal feet. No clear division is
discernible between pigmented and non-pigmented forms since specimens from all
localities show the above pattern, but one separate and large collection from
Christiansborg shore (15.L49) is completely without colour. The outside edge of
the filaments is thickened in places suggesting rudimentary stylodes, but they are
much less well developed than in the genus Dasychone. The branchial membrane
reaches along one-sixth of the length of the branchial filaments ; the two palps are
foliaceous and a little longer than the membrane.
There are 4-5 thoracic chaetigers and about 100 abdominal segments. Neuropodial
uncini appear at the 2nd foot. Thoracic capillaries are of two kinds, short with a
prominent limbate termination (not spatulate) and long with a slightly limbate end ;
thoracic uncini are short, avicular. Abdominal capillaries and uncini are identical
with thoracic bristles.
Augener (1918) placed this species in the genus Demonax Kinberg, which, however,
has pickaxe-shaped chaetae in the thoracic neuropodia ; Monro, (1930) placed it
in the genus Euratella Chamberlin, which has no collar and only one type of thoracic
capillary. It is here considered that the species is referable to Laonome.
Genus DASYCHONE Sars, G.O., 1861
Both branchial lobes equal, not in the form of a spiral. Stylodes on the branchial
filaments. Prominent collar. Pickaxe-shaped chaetae absent.
Dasychone lucullana (Delle Chiaje), 1828
Dasychone lucullana, Fauvel, 1927a, pp. 320-321, fig. no, m-s.
Localities. Off Accra: Stn. 35 (1), 8 mm. long; Stn. 73 (1), 15 mm. long;
Stn. 131 (1), 16 mm. long, all specimens complete.
The two branchial lobes are each made up of 13-14 filaments which carry slender,
colourless stylodes dorsally. Along the filaments there are numerous violet pigment
spots, which, when the branchiae are closed, give the effect of successive girdles of
colour along the lobes. A small branchial membrane unites the bases of the gills ;
there are two prominent canaliculate palps.
The body, other than the branchiate portion, is uniformly brown, with a small
violet pigment spot between the parapodial rami. There are 8 thoracic and between
30 and 40 abdominal segments. Thoracic uncini which appear at the 2nd chaetiger,
are avicular with slight extension at the base ; thoracic capillaries are limbate,
short and long. Chaetae in the abdominal region are similar to thoracic types except
that long capillaries, only slightly limbate, appear in far posterior segments.
Distribution. Mediterranean and Adriatic Seas and from Senegal.
THE POLYCHAETE FAUNA OF THE GOLD COAST 141
Genus POTAMILLA Malmgren, 1865
Branchial lobes symmetrical, not in a spiral, without stylodes. With a prominent,
lobed collar. Thoracic notopodial capillaries limbate and spatulate, neuropodial
chaetae avicular uncini and pickaxe-shaped. Abdominal notopodial uncini avicular
and neuropodial capillaries limbate.
Potamilla casamancensis Fauvel, 1902
Potamilla casamancensis Fauvel, 1902, pp. 101-105, fig. 46-55, from the estuary of the River
Casamance, Senegal.
Localities. Off Accra : Stn. 12 (1), complete, 16 mm. long ; Buchanan Survey
in 11 metres off Lighthouse (2), complete, 14 and 24 mm. long ; Buchanan Survey
in 5-5 metres off the Lagoon (1), incomplete, 8 mm. long.
There is an indentation in the collar dorsally, forming a deep groove which con-
tinues down the dorsal surface of the thorax ; ventrally there is only a very slight
break in the anterior border of the collar.
Ten to sixteen filaments make up each branchial lobe ; the filaments are without
eye-spots. There are 5-7 thoracic chaetigers ; uncini and pickaxe-shaped chaetae
appear at the second. The limbate capillaries on the first segment are in a longi-
tudinal row, thereafter they are lateral. Spatulate capillaries appear at the second
chaetiger and are normally few in number. Limbate capillaries are present in all
segments, both thoracic and abdominal ; spatulate chaetae have not been seen in
the abdomen. Uncini in the thorax are avicular with long shafts ; in the abdomen
they are short.
Distribution. P. casamancensis is only known through the above records.
Family Serpulidae
Body divided into two regions ; [a) thorax, with few segments, having notopodial
capillaries and neuropodial uncini, and (b) abdomen, with numerous segments,
having notopodial uncini and neuropodial capillaries. First segment with a collar.
With a thoracic membrane. Branchiae form a funnel around the mouth. Usually
with an operculum. Tube calcareous.
Key to Genera
1. Body symmetrical ............ 2.
Body assymetrical. Tube spirally coiled ....... Spirorbis.
2. Without an operculum .......... Salmacina.
With an operculum ............. 3.
3. Chaetae of the first chaetiger bayonet-shaped with two basal stumps . . . 4.
Chaetae of the first chaetiger without basal stumps . . . . . . 5.
4. Operculum simple, funnel-shaped ........ Serpula.
Operculum compound, with a central crown of spikes ..... Hydroides.
5. Operculum globular, transparent ........ Apomatus.
Operculum horny, with a cylindrical or conical cap , Vermiliopsis.
I 4 2
THE POLYCHAETE FAUNA OF THE GOLD COAST
Genus SALMACINA Claparede, 1868
Without an operculum and with few branchiae which are frequently bent distally.
Uncini pectinate with a large inferior tooth.
Salmacina incrustans Claparede, 1868
Salmacina incrustans, Fauvel, 1927a, pp. 378-380, fig. 129, /.
Localities. Numerous specimens were collected from : (1) Tenpobo shore, on
rocks, 4-ii.5o ; (2) Winneba rock face, sheltered side, 22.xi.49, forming compact
masses about the Polyzoan, Waterispora cucullata ; (3) Apam shore, 16.ii.49 ; (4)
Axim hospital reef, 13.lv. 49, L.W.M., some specimens were present on a tube of
Sabellaria spinulosa var. alcocki.
The majority of these specimens measure between 2 and 3 mm. Tubes are white,
cylindrical in shape, and very fragile.
Fig. 30. Chaetae of (a) Potamilla casamancensis , pickaxe-shaped bristle from a thoracic
neuropodium, x 350 (stn. 49) ; (b) Salmacina incrustans, a notopodial chaeta from
the first thoracic segment x 200, (Winneba) ; (c) and (d) Hydroides arnoldi, thoracic
and abdominal uncini respectively, x 400 (e), (/) and {g) Apomatus similis : (e)
A pornatus -type chaeta from a posterior thoracic segment, x 100, (/) limbate chaeta
from a thoracic segment, (g) scythe-like chaeta from a median abdominal segment
X 150; (h) Serpula vermicularis , trumpet-shaped abdominal chaeta, x 200 (Stn. 115) ;
(;') Vermiliopsis multicristata, geniculate abdominal chaeta, x 200.
There are four pairs of gill filaments and seven thoracic chaetigers. A notopodial
chaeta from the first of these is shown in Text-fig. 30, b. This form is characteristic
for the species.
Distribution. Mediterranean Sea and Atlantic Ocean.
THE POLYCHAETE FAUNA OF THE GOLD COAST 143
Genus HYDROIDES Gunnerus, 1768
Chaetae of the first chaetiger bayonet-shaped, with two humps at the base of the
blade. Thoracic chaetae limbate ; abdominal chaetae trumpet shaped. Uncini in
thorax and abdomen with coarse teeth. Operculum funnel-shaped, with radii
ending bluntly or in fine points, and with a crown of horny spines rising from its
centre.
Pixell (1913) suggested that Hydroides should include those species with opercular
spines having lateral processes and that species devoid of these lateral processes
should be included within the genus EupomcUus. This division is not adopted in this
paper.
Key to Species
1. Opercular spines all alike ........... 2.
Opercular spines not all alike ........ H. spinosus.
2. Opercular spines with lateral processes . . . . . . H. norvegica.
Opercular spines without lateral processes . . . . . . . H. arnoldi.
Hydroides norvegica Gunnerus, 1768
Hydroides norvegica, Fauvel, 1927a, pp. 356-7, fig. 122, i-o.
Locality. Scraped off copper sheathing of boat and body, Gold Coast, (50).
The species is immediately recognizable by its opercular structures. The basal
disc is soft and white, bearing marginally between 20 and 30 crenulations. Fre-
quently the distal horny disc is missing but, when present, it carries 13 to 15 large
spines with smaller lateral ones. A small vertical spine is situated in the centre of
the opercular disc. The operculum itself is a modified branchial filament, the ventral-
most of one side or the other ; whichever it may be, and there is no constancy, the
other is always broken off.
Chaetae of the first of the seven thoracic chaetigers have two humps below the
terminal point and below the humps there are a few fine denticulations. Thoracic
uncini have seven teeth and abdominal uncini five.
Distribution. Cosmopolitan.
Hydroides spinosus (Pixell), 1913
Eupomatus spinosus Pixell, 1913, p. 78, pi. 8, fig. 5.
Localities, (i) Tenpobo, No. 1 Reef, 21.xi.49 (1), complete, 15 mm. long.
(2) Off Accra : (a) Trawled 2 miles west of Densu River, 2. hi. 49 (1), incomplete,
10 mm. long ; (b) Stn 99 (3), one complete, 10 mm. long, on tubes of Turritella
annulata Kiener ; Stn. 127 (1), complete, 11 mm. long, on a tube of T. annulata.
These specimens differ only slightly from the original description (Pixell, 1913).
Thus the basal disc has frequently more than 32 long, pointed teeth and the number
of tall strong spines on the upper disc, is not constantly 11 but varies, some speci-
mens having 8, 9 or 10. The shape of all the spines except one is identical, they each
have a terminal hook pointing outwards, a near-terminal one pointing down wards
i 4 4 THE POLYCHAETE FAUNA OF THE GOLD COAST
and inwards and an inner median near the base. The exceptional one is longer than
the others, curves over them, and has only a basal hook.
Distribution. H. spinosus has previously been recorded from the Suez Canal
area and the Cape of Good Hope.
Hydroides arnoldi Augener, 191 8
Hydroides arnoldi Augener, 1918, pp. 595-598, text-fig. 107, pi. 6, figs. 151, 152, from Togo,
Dahomey, and the Island of Annobon.
Locality. Axim shore, 14.lv. 49 (3), incomplete, 4-11 mm. long.
There are seven opercular spines which have an inner basal hook but are devoid
of other processes. The largest, and possibly oldest, specimen has almost straight
spines with blunt ends (Augener, 1918, pi. 6, fig. 152), but the smallest, and possibly
youngest, has spines ending in a curved tip. It is suggested that normal wear on
the operculum has produced these different shapes and the species may be expected
to vary in this character even more than is noted here.
Thoracic and abdominal uncini are illustrated in Text-fig. 30, c and d.
Distribution. H. arnoldi is only known from the above records.
Genus APOMATUS Philippi, 1844
With a globular transparent operculum. Branchial filaments with eye-spots and
short connective membrane. No special notopodial chaetae on the first thoracic
segment, which is without neuropodia, but with characteristic Apomatus-type
bristles on the last segments (Text-fig. 30, e).
Apomatus similis Marion and Bobretzky, 1875
Apomatus similis, Fauvel, 1927a, pp. 385-387, fig. 131 k—p.
Locality Off Accra, Stn. 115 (10), with tubes, on the Polyzoan Steganoporella
buskii Harmer. The specimens measure up to 6 mm. in length.
The branchial lobes are made up of 10 to 16 filaments each. The second dorsal
filament carries a globular transparent operculum distally, but it is otherwise un-
modified and combines a branchiate and operculate function. This represents a
primitive stage in the modification of a branchial filament to an operculum. Com-
plete modification is found in the majority of Serpulidae, but there is no trace of it
in Filograna and Protula.
There are seven thoracic and well over 60 abdominal segments. The majority of
thoracic chaetae are limbate (Text-fig. 30, /), but in the last 3-4 segments of the
thorax characteristic Apomatus-type bristles appear (Text-fig. 30, e). Anterior
abdominal segments are achaetous. Neuropodia in the middle of the abdomen have
very small scythe-like chaetae (Text-fig. 30, g), but in four posterior segments long
capillaries are present. Abdominal and thoracic uncini are alike, with numerous
small teeth and a long basal one.
Tubes are white and semi-cylindrical, with slight lateral crenulations.
Distribution. English Channel, Mediterranean Sea, Atlantic Ocean,
THE POLYCHAETE FAUNA OF THE GOLD COAST 145
Genus SERPULA Linnaeus, 1758
Chaetae of the first thoracic segment bayonet-shaped, with two humps at the
base of the blade. Operculum funnel-shaped with radii ending bluntly along the
margin. Thoracic chaetae limbate, abdominal chaetae trumpet-shaped. Uncini
with few, but stout teeth.
Serpula vermicularis Linnaeus, 1767
Sevpula vermicularis, Fauvel 1927a, pp. 351-352, fig. 120, a-q.
Localities, (i) Tenpobo reef, 12.xi.49 (2), incomplete, 10 and 20 mm. in length,
without tubes. (2) Off Accra, Stn. 115 (3), complete, up to 20 mm. in length.
These specimens appear to be smaller than is normal for the species (Fauvel,
1927a), but those from Stn. 115 were taken from tubes completely attached to the
polyzoan, Steganoporella buskii, none of the tubes growing free, and they may be
juveniles. The tubes are pink in colour, and triangular in shape with a prominent
dorsal keel.
There are 16-20 branchial filaments in each lobe united by a short basal membrane.
The first dorsal filament of one side is completely adapted as an operculum, with a
funnel-shaped termination having a crenulated surface. The first dorsal filament on
the opposite side to that bearing the operculum is partially modified ; it is similar
to the opercular filament, but is smaller and lacks the funnel-like termination.
There are seven thoracic chaetigers the first without neuropodia. Apart from the
bayonet-shaped chaetae with two basal humps on the first thoracic segment,
notopodial-chaetae are unmodified capillaries. Abdominal neuropodial chaetae are
trumpet -shaped (Text-fig. 30, h).
Distribution. Cosmopolitan.
Genus VERMILIOPSIS St. Joseph, 1906
Operculum a horny cylindrical or conical cap. No special chaetae on the first
chaetiger, which is without neuropodia. Apomatus-type chaetae in posterior thoracic
segments. Abdominal chaetae geniculate (Text-fig. 30, j).
Key to Species
1. With seven thoracic chaetigers ........ V. multicristata.
2. With nine thoracic chaetigers ....... V. prampramiana.
Vermiliopsis multicristata (Philippi), 1844
Vermiliopsis multicristata, -Fauvel, 1927a, pp. 365-366, fig. 125, k-s.
Locality. Off Accra, Stn. 115 (6), some complete, up to 10 mm. long, taken from
tubes attached to the polyzoan Steganoporella buskii.
This species is characterized by its tube bearing 5-6 longitudinal ridges. There
are about 10 branchial filaments to each lobe. The operculum is horny and cylindrical
146
THE POLYCHAETE FAUNA OF THE GOLD COAST
to conical in shape. In the thorax there are seven chaetigers and in the abdomen
upwards of 50. The majority of thoracic chaetae are limbate, but in posterior
segments a few Apomatus-type bristles appear. Abdominal chaetae are short and
geniculate (Text-fig. 30, j) anteriorly, but posteriorly long capillaries appear.
Distribution. Mediterranean Sea and Atlantic Ocean — Madeira and the Azores.
Vermiliopsis prampramiana Augener, 1918
Vevmiliopsis prampramiana Augener, 1918, pp. 603-604, pi. 7, fig. 256, from Pram Pram.
This species is only known through this record, and is the only species of Vermiliop-
sis known with nine thoracic chaetigers. Augener's description is otherwise in-
complete, and it is difficult to assess the species relationship with other members
of the genus or family.
Genus SPIRORBIS Daudin, 1800
Body assymetrical, with less than five thoracic segments. Opercular peduncle
without pinnules. Tubes spirally coiled, dextral or sinistral.
4-
Key to Sub-genera
Tube dextral ........ ... 2
Tube sinistral .......
3
With 4 thoracic segments .....
(Paradexiospira)
With 3 thoracic segments .....
(Dexiospira)
With 4 thoracic segments .....
(Paralaeospira)
With 3 thoracic segments .....
4
Chaetae of the first thoracic chaetiger limbate
(Leodora)
Chaetae of the first chaetiger with a crenulated wing
. (Laespira)
This genus was, surprisingly, absent from the collections, but it is almost certainly
present in the Fauna, and, for this reason the above key has been included, (from
Fauvel, 19270, pp. 389-391). Augener (1918, pp. 607-608), reported Spirorbis sp.
from the " Cape Coast Castle", probably Christiansborg Castle, but his description
does not permit further identification. When, and if, species are collected further
reference to Fauvel should be made.
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Augener, H. 1913. Polychaeta Errantia. Die Fauna Sudwest-Austt -aliens, 4 : 65-304,
2 pis., 42 text-figs.
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1924. Polychaeta von Neuseeland. I, Errantia. Medd. naturh. Foren. Kobenhaven,
75 : 241-441, 11 figs.
THE POLYCHAETE FAUNA OF THE GOLD COAST 147
Baird, W. 1864. Description of a new species of Annelide belonging to the family Amphino-
midae. Trans. Linn. Soc. London, 24 : 449-450, pi. 45.
Bergstrom, E. 1914. Zur Systematik der Polychaeten familie Phyllodociden. Zool.
Bidrag. Uppsala, 3 : 37-224, 81 text-figs.
Chamberlin, R. V. 1919. The Annelida Polychaeta. Mem. Mus. Comp. zool. Harvard,
48 : 1-5 14, pis. 1-80.
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Crossland, C. 1903. On the marine fauna of Zanzibar and British East Africa, from collec-
tions made by Cyril Crossland in the years 1901 and 1902. Polychaeta, Pt. 2, Eunicidae.
Proc. Zool. Soc. London, 2 : 129-144, pis. 14 and 15, text-figs. 12-15.
1904. The marine fauna of Zanzibar and British East Africa from collections made by
Cyril Crossland in the years 1901 and 1902. Polychaeta, Pt. 3. Ibid., 1 : 287-330,
pis. 20 to 22, text-figs. 43-66.
1924. Polychaeta of Tropical East Africa, the Red Sea and Cape Verde Islands collected
by Cyril Crossland, and of the Maldive Archipelago collected by Professor Stanley Gardiner,
M.A., F.R.S. Ibid. Pt. 1 : 1-106, text-figs. 1-126.
Day, J. H. 1934. O n a collection of South African Polychaeta, with a Catalogue of the
species recorded from South Africa, Angola, Mosambique and Madagascar. J. Linn.
Soc. London, Zool. 39 : 15-82, 16 text-figs.
1949. Polychaeta of St. Helena. Ibid. 41 : 434-451, 4 text-figs.
1 95 1. The Polychaet Fauna of South Africa. Pt. 1 : The Intertidal and Estuarine
Polychaeta of Natal and Mozambique. Ann. Natal Mus. 12 : 1-67, 8 text-figs.
1953- The Polychaet Fauna of South Africa. Pt. 2 : Errant Species from Cape Shores
and Estuaries. Ibid. 12 : 397-441, 1 map. 6 text-figs.
Eiilers, E. 1887. Reports of the annelids of the dredging expeditions of the U.S. coast
survey steamer "Blake." Mem. Mus. Comp. Zool. Harvard. 15 : 1-335, °° plates.
Fauvel, P. 1902. Ann61ides Polyene tes de la Casamance (Senegal, Africa). Bull. Soc.
Linn. Normandie, 5th series, 5 : 59-105, figs. 1-55.
191 1. Annelides Polychetes du Golfe Persique. Arch. Zool. Exper. gen. Paris. (5),
6 : 353-439, pis. XIX-XXI.
1914- Annelides Polychetes de San Thome (Golfe de Guinee). Ibid. 54: 105-155, pis.
VII-VIII.
1 91 9. Annelides Polychetes de Madagascar, de Djibouti, et du Golfe Persique. Ibid.
58 : 315-473, Pis- 15-17-
1923. Polychetes errantes. Faune de France, Paris 5 : 488, 1-181 text-figs.
1927a. Polychetes sedentaires. Ibid. 16 : 1-494, 152 text-figs.
19276. Contribution a l'etude de Faune du Cameroun. Polychaeta. Faun. Colon.
franc, 1 : 5^3-533, 2 figs.
1932. Annelida Polychaeta of the Indian Museum, Calcutta. Mem. Indian Museum
Calcutta 12 : 1-262, 9 pis., 40 text-figs.
1933- Annelides Polychetes du Golfe du Pei Tcheu Ly de la collection du Musee Hoang
ho Pai ho. Publ. Mus. Hoang. ho Pai do de Tien Tsin, 15 : 1-67, 6 text-figs.
1936. Remarques sur les Nereidiens, Nereis succinea Leuckart et Nereis lamellosa Ehlers.
Bull. Soc. Zool. France, 61 : 307-314.
1950. Contribution a la Faune des Annelides Polychetes du S6negal. Bull. Inst, francais
d'Afrique noire, 12 (2) : pp. 335-394.
Fyfe, M. L. 1952. List of New Zealand Polychaetes. Bull. New Zealand Dept. Sci. Industrial
Research, 105 : 1—38.
Gravier, C. 1909. Ann61ides polychetes recueillis a Payta (Perou). Arch. zool. exp. gen.
Paris, 10 : 617-659.
Hartman, O. 1939. Polychaetous Annelids. Pt. 1 : Aphroditidae to Pisionidae. Allan
Hancock Pacific Exped. 7 : 1-156, 28 pis.
1944a. Polychaetous Annelids. Pt. 5 : Eunicea. Ibid. 10 : 1-238, 18 plates.
zool. 3, 2. 12
148 THE POLYCHAETE FAUNA OF THE GOLD COAST
Hartman, O. 19446. Polychaetous Annelids, Part 6, Ibid., 10 : 311-389, pis. 27-42.
1948. The Marine Annelids erected by Kinberg with notes on some other Types in the
Swedish State Museum. Ark. Zool. Stockholm, Band 42A, no. 1 : 1-137, pis. 1-18.
1950. Polychaetous Annelids, Goniadidae, Glyceridae, Nephtyidae. Allan Hancock
Pacific Expd. 15 : 1-181, 19 plates, 3 text-figs.
1 95 1. The Littoral Marine Annelids of the Gulf of Mexico. Publ. Inst. Marine Sci.
Univ. Texas 2 (1) : 7-124, 27 pis.
Kinberg, J. 1856. Nya slagen och arter af Annelider. Fork. Oefv. Vet. Akad. Stockholm
12 : 381-388.
1857-1910. Kongliga Svenska Fregatten Eugenics Resa omkring Jovden under C. A.
Virgin 1851-53. Zool. 3, Annulater, Uppsala-Stockholm, 78 pp., 29 plates.
McIntosh, W. C. 1885. Report on the Annelida Polychaeta collected by H.M.S. " Chal-
lenger " during the years 1873-6. " Challenger " Rep. Zoo. 12 : 1-554, pl s - I_ 55 an d
ia-39a.
1903. Marine Annelids (Polychaeta) of South Africa, I. Marine Investigations in South
Africa, 3 : 17-56, 4 pis.
1924. Preliminary note on a collection of South African polychaetes. Report Union
South Africa Fish Mar. Biol. Survey, Cape Town, 3, No. 2, 1 p.
Monro. C. C. A. 1930. Polychaeta Worms. " Discovery " Reports, 2 : 1-222, 91 text-figs.
1933- The Polychaeta Errantia collected by Dr. C. Crossland at Colon, in the Panama
Region, and the Galapagos Islands during the Expedition of the S.Y. " St. George ".
Proc. Zool. Soc. London, 1933, (1) : 1-96, 36 text-figs.
1936a. Polychaete Worms 2. " Discovery " Reports, 12 : 59-198, 34 figs.
19366. Notes on some Polychaeta from the Congo Coast. Rev. Zool. Bot. Afr. 28 : 245-
248, 1 fig.
Pixell, H. L. M. 191 3. Polychaeta of the Indian Ocean, together with some species from
the Cape Verde Islands. The Serpulidae. Trans. Linn. Soc. London, 2nd. Ser., Zoology,
16 : 69-92, pis. 8 and 9.
Pruvot, G. 1930. Annelides polyene tes de Nouvelle-Caledonie recueillies par M. Francois.
Arch. Zool. exp. gen. Paris, 70 : 1-94, 8 figs., 3 pis.
Quatrefages, M. A. de. 1 865. Histoire des Anneles, 1, Paris, 588 pp.
Takahasi, K. 1938. Polychaeta, collected by the " Misago " during the Zoological Survey
around the Izu Peninsula. Sci. Rep. Tokyo Bunrika Daigaku, Sec. B, 3 (57) : 192-220,
1 pi.
Treadwell, A. L. 1900. The polychaetous Annelids of Porto Rico. Bull. U.S. Fish Comm.
Washington, 20 : 18 1-2 10, 81 figs.
Willey, A. 1904. Littoral Polychaeta from the Cape of Good Hope. Trans. Linn. Soc.
London. Zool. 9 (2) : 255-268, pis. 13-14.
1905. In Herdman. Roy. Soc. Report on Pearl Oyster Fisheries, Ceylon. Pt. 4 : 243-
324, pis. 1-8.
2 SEP 1955
PRINTED IN GREAT BRITAIN BY
ADLARD AND SON, LIMITED
BARTHOLOMEW PRESS. DORKING.
2 JUL 1955.
A REVISION OF THE OCTOPODINAE
IN THE COLLECTIONS OF THE
BRITISH MUSEUM
GRACE E. PICKFORD
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 3 No. 3
LONDON : 1955
A REVISION OF THE OCTOPODINAE IN THE
COLLECTIONS OF THE BRITISH MUSEUM
BY
GRACE E. PICKFORD
(Bingham Oceanographic Laboratory, Yale University)
Pp. I51-167.
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 3 No. 3
LONDON: 1955
THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, is
issued in five series corresponding to the Departments
of the Museum, and an Historical Series.
Parts will appear at irregular intervals as they become
ready. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.
This paper is Vol. 3, No. 3 of the Zoological series.
PRINTED BY ORDER OF THE TRUSTEES OF
THE BRITISH MUSEUM
Issued June, 1955 Price Five Shillings
Part L— FURTHER LIGHT ON THE OCTOPUS RUGOSUS
PROBLEM
By GRACE E. PICKFORD
In spite of the efforts of Robson (1929), which form a landmark in the history of
octopodan taxonomy, the Octopus rugosus problem remains unsolved, an outstanding
source of confusion obscuring the correct identification of small, rough, vulgaris-like
specimens from all parts of the world. Some years ago the writer reviewed the western
Atlantic forms which had been, or could be, assigned to Octopus rugosus. It was
possible to show that no such species can be found in American waters and that
western Atlantic specimens previously referred to it were either 0. vulgaris or,
occasionally, belonged to one of the other Caribbean species (Pickford, 1945, 1946).
It is believed that the problem can best be solved by a zoogeographical approach,
pari passu with increasing knowledge of each local fauna to the point that makes
the recognition of its component species a matter of certainty. Through the work
of Voss (1950, 1 95 1, etc.) and the author we have now arrived at this state of know-
ledge of the American east coast fauna. However, the situation in other parts of
the world is less satisfactory. Octopodan taxonomy has always been hampered by
the lack of adequate series of specimens collected from any one locality and this,
together with the confusing effects of different states of preservation, has made
recognition of the limits of variation within a single species all but impossible.
In the present contribution an attempt has been made to clarify the status of
those specimens from the eastern Atlantic and South Africa which Robson identified
as 0. rugosus and which are now in the collections of the British Museum. A survey
of bodily characteristics, irrespective of species assignation, is given in Table I. It
will be seen that only 7 of the 22 specimens are sexually mature, 4 are well grown
but immature, and n are juvenile. In a difficult problem of this nature it is
obvious that no definite conclusions can be based on juvenile, nor even on sexually
immature specimens, except in so far as they can reasonably be assigned to adult
forms with well established characteristics.
Octopus rugosus, as it is generally understood, is a species closely resembling 0.
vulgaris but smaller in size ; the head is broader, the arms shorter, and the rough,
finely granulated skin is darkly reticulated with black. None of these characters,
taken singly, can be used with any degree of certainty for the separation of the two
species. Taken together the resulting picture is indistinguishable from that of
juvenile and immature specimens of 0. vulgaris, so that it may be conjectured that
0. rugosus is merely the young of 0. vulgaris (Pickford, 1945). This point of view
is that taken by the writer who believes that it is necessary to prove that an Atlantic
specimen assigned to 0. rugosus is not a young 0. vulgaris before it can be considered
to belong to any other species ; a rugosus-like specimen should be assigned to 0.
zool. 3, 3. 13
152 REVISION OF OCTOPODINAE IN THE BRITISH MUSEUM
vulgaris unless there is conclusive evidence that it does not belong to this species.
Robson, on the other hand, used the name 0. rugosus as a hold-all to receive those
vulgaris-like specimens about whose identification he was uncertain and, as a result
of this policy, it is not surprising to find that the majority of his eastern Atlantic
specimens, as indicated above, are juvenile or immature.
We may approach the problem from another angle. Bosc's Sepia rugosa came
from Senegal and the type is lost. Lamarck (1799) gives no locality for the two
specimens to which he gave the name 0. granulatus. He points out that this species
may be merely a variety of 0. vulgaris and concludes that it is probably the same
as Bosc's rugosus. The types are lost and, in the absence of evidence to the contrary,
it appears advisable to relegate the name 0. granulatus to the synonymy of 0.
vulgaris. A similar conclusion is probably justified in the case of 0. tuber culatus
Blainville, the type of which is also lost. However, since the species is from Sicily,
its status should be re-examined in connexion with a more thorough taxonomic
study of the Mediterranean population.
It must be pointed out that the original specimens of 0. rugosus and 0. tuber cu-
latus, not to mention numerous other specimens referred by various authors to these
species or to 0. granulatus, were reported from geographical areas in which 0.
vulgaris is known to occur. Therefore, 0. rugosus (to confine the discussion to one
of the three problematical forms) is not a geographical race or subspecies of 0.
vulgaris. Either it is a distinct species or it is inseparable morphologically from the
extremes of the 0. vulgaris population. This is an important consideration. If two
species occur together in the same geographical area they must form two separate
breeding populations.
The search for 0. rugosus is therefore the search for a West African vulgaris-like
sibling species which is distinct as. a breeding population from 0. vulgaris with
which it is sympatric. As the result of previous investigations (Pickford, 1945, 1946)
it can definitely be stated that 0. rugosus does not occur in the western Atlantic
but the problem of Indo-Pacific forms that have been assigned to this species awaits
further clarification. However, if the existence of a true 0. rugosus cannot be
established in the eastern Atlantic it is highly improbable that anything recorded
from Indo-Pacific waters could legitimately be assigned to this species.
The study of the British Museum specimens reviewed below leads to the following
conclusion :
(1) Nearly all specimens from the eastern Atlantic are probably or certainly
referable to 0. vulgaris. It is probable that the correct assignation of the juvenile
specimens will always remain in doubt but 6 of the 7 adults present no features
which would permit their separation from 0. vulgaris.
(2) Among the series there is one juvenile specimen from Madeira that has a
distinctive pattern of zebra bands on the arms. Although the characteristics (Table I,
B.M., 1898. 5. 10.2) are similar to those of 0. vulgaris the correct assignation of this
specimen must remain in doubt.
(3) One specimen from the Cape Verde Islands (1889. 4. 24. 17), a mature male,
certainly represents a distinct species on account of the length of the ligula which
is quite outside the range for 0. vulgaris. A similar male and also a female that
REVISION OF OCTOPODINAE IN THE BRITISH MUSEUM 153
apparently belong to the same species have been described by Adam (1952) from
the Cape Verde Islands as Octopus sp. The question arises whether these individuals
should be arbitrarily assigned to 0. rugosus, with which they show some general
agreement, or whether it would be preferable to designate them by a new name.
The original description of 0. rugosus does not permit recognition with any degree
of certainty. Bosc's figures (1792, Figs. 1 and 2), if correctly drawn, show an animal
of 42 mm. mantle-length with moderately long arms (MAI 32) and an unusually
shallow web (WDI 13-5) ; the suckers are of moderate size (SnI 9 • 5) but none are
specially enlarged. In favour of assigning the Cape Verde specimens to this species
is the geographical area from which it came, as well as the general appearance and
bodily proportions, except for the shallow web. The only contrary evidence is the
shallow web and the uncertainty whether Bosc's 0. rugosus was not, after all, just
a young vulgaris. For the sake of clarity and to avoid the confusion that might
result from the use of a name which is, by now, so clouded in obscurity, it has
seemed preferable to propose a new name for this Cape Verde species.
(4) The problem of the specific affiliation of the South African rugosus-like popu-
lation remains unsettled. The individuals that have been examined, from the Cape
Province and Natal, appear to belong to the same species and to represent a moderately
uniform group. Robson admits (1929, p. 59) that true 0. vulgaris occurs in South
African waters. The two males, from Port Elizabeth and from Natal respectively,
have the characteristic minute ligula of 0. vulgaris}- Massy (1925) found the same
feature in males which she identified as 0. granulatus in the collections of the Natal
Museum (LLI 1-7 and 1-65 respectively, calculated from her measurements).
There is a small but possibly significant detail regarding the repeated reversal of
the direction of coiling of the horn in the spermatophore of one of the males (B.M.,
1890 .9.4.4), but our information concerning the range of variation in spermatophores
of 0. vulgaris is inadequate to assess this feature. Also the females, with ripe ovaries,
are unusually small for sexually mature females of 0. vulgaris ; but this again
amounts to no more than a suggestion. Possibly the South African forms represent
a geographical race or subspecies of 0. vulgaris with these slightly different features.
Such a conclusion could only be confirmed by the statistical analysis of a long series
of specimens.
A detailed account of the specimens follows :
DISCUSSION OF SPECIMENS WHICH ARE PROBABLY OR CERTAINLY
REFERABLE TO OCTOPUS VULGARIS LAM.
English Channel
(1) B.M., 1927. 2. 10. 2. Octopus rugosus (Bosc), Robson (1929). Plymouth. C.92.
It was pointed out in a previous paper (Pickford, 1946) that this specimen is
clearly a female of 0. vulgaris. I take this opportunity to publish the characteristics
listed in Table I and to call attention to the following features : the arms are un-
1 Attention must again be called to Robson's mistake in tabulating the ligula-length indices for
O. vulgaris. It appears that in this one case he referred the length of the ligula to the mantle length
instead of to the length of the hectocotylised arm, thus giving misleadingly high values (see Pickford,
1945)-
154
REVISION OF OCTOPODINAE IN THE BRITISH MUSEUM
usually short and the web rather deep but the suckers and gills are normal ; there
is a very conspicuous lateral seam round the mantle, no doubt an accident of pre-
servation ; the ovary is quite small and the ova are minute.
Table I. — Characteristics of specimens from the eastern Atlantic, and South Africa
which were referred by Robson (1929) to the species Octopus rugosus (Bosc), in the
collections of the British Museum*
B.M. No. Sex ML MWI HWI MAI WDI SnI Gill
English Channel
1927.2. 10.2
$ imm.
70
79
5i
49
39
14-3
11
Mediterranean
1867.3.27.9
6* juv.
21
67
72
35
15
8-6
8-5
Madeira
1898.5. 10.2
$ imm.
32
78
69
27
21
II-O
ca. 8
1912.12.31.118
juv.
8
1912.12.31.119
juv.
6.5
Canary Islands
1954.2.25.1
juv.
8-5
—
—
—
—
—
—
1954.2.25.2
juv.
8
—
—
—
—
—
—
1954.2.25.3
juv.
6
Cape Verde Islands
1851.1.24.1
$ imm.
50
84
70
28
24
16
8
1855.9.29.2
6* juv.
31
68
56
3i
21
9.7
ca.8
1889.4.24. 17
6* mat.
24
92
7i
40
28
8-3
9
Ascension
1889.4.24. 14
$ imm.
57
88
73
32
28
14
ca. 10
(55)t
(9i)
(73)
(24)
—
(12-7)
—
West Africa
1928.3. 17. 1
£ mat.
24
58
63
21
21
16-7
ca. 6-'
1928.3. 17.2
juv.
13
46
62
—
—
"*5
—
1928.3. 17.3
juv.
15
93
—
—
—
15-3
—
1889.4.24. 15
juv.
7*5
South Africa
1890.9.4.4
c£ mat.
30
77
74
27
23
15
8-5
1924.9.9. 11
$ mat.
41
1924.9.9. 12
$ mat.
45
1924.9.9-13
$ mat.
59
1926.10.20.30
$ mat.
27
(28) J
(86)
—
(34*5)
—
(10-7)
—
1889.4.24. 18
juv.
12
* Two females from the " South Coast " could not be found (B.M., 1872 .2.3. 12-13). Three specimens
(1865.5.23. 1) from an unknown locality omitted. Data based on new measurements unless otherwise
indicated. Symbols as in Pickford (1945).
t Data from Hoyle (1886).
I Data from Massy (1927).
REVISION OF OCTOPODINAE IN THE BRITISH MUSEUM 155
Mediterranean
(2) B.M., 1867.3.27.9. Octopus rugosus (Bosc), Robson (1929). Mediterranean,
purch. of Mr. Cutter. C.104.
This is a very young male, not a female as listed by Robson. As far as can be
determined it is probably referable to 0. vulgaris.
Madeira
(3) B.M., 1898.5. 10.2. Octopus rugosus (Bosc), Robson (1929). Porto Santo,
Madeira, H. S. Wellcome Esq. C.86.
This specimen, an immature female, is remarkable for its coloration. There are
light and dark zebra bands on the arms and it is very dark and blotchy on the
back and head. The ovary is undeveloped but has moved to a ventral position and
is possibly rather advanced for a vulgaris of this size ; the ova are minute. The
jaws have been dissected out and the radula is mounted on a slide. The bodily
proportions are similar to those of a typical vulagris but on account of the peculiar
coloration I hesitate to refer the animal to this species. It will be necessary to
await further collections from this region to establish the existence of a form with
banded arms and, until such time, it may be considered to be merely an unusual
specimen of 0. vulgaris.
(4) B.M., 1912.12.31.118 and 119. Octopus rugosus (Bosc), Robson (1929). Pedro
de Malha, Porto Santo, Madeira, 60 fathoms, R. Kirkpatrick Esq. C. 152-153.
Two very young specimens with the characteristic rhomb of 4 papillae on the
back commonly seen in juvenile specimens of 0. vulgaris ; probably correctly
referred to this species.
Canary Islands
(5) B.M., 1954. 2. 25. 1-3. Octopus rugosus (Bosc), Robson (1929). Canaries, R.
McAndrew. C. 100-102.
Three very young specimens, probably all referable to 0. vulgaris. The two
larger specimens each have the characteristic rhomb on the back.
Cape Verde Islands
(6) B.M., 1851.1.24.1. Octopus rugosus (Bosc), Robson (1929). St. Jagos, C. de
Verd, Ch. Darwin Esq., No. 122. C. 85.
An immature female, very dark, mottled and rough. The ovary is small and
apical ; the ova minute. The jaws have been dissected out. The suckers are large,
as in vulgaris, the arms moderately long and the web moderately deep. There seems
to be no reason why this specimen should not be assigned to 0. vulgaris.
(7) B.M., 1855.9.29.2. Octopus rugosus (Bosc), Robson (1929). St. Vincent, Mr.
MacGillivray coll. C. 119.
156 REVISION OF OCTOPODINAE IN THE BRITISH MUSEUM
This specimen is an immature male. The head is rather warty, the body smooth.
The total absence of a hectocotylus at a mantle-length of 31 mm. indicates that it
belongs to a large species and there seems to be no reason why it should not be
assigned to 0. vulgaris.
Ascension
(8) B.M., 1889. .4.24.14 Octopus occidentalis Hoyle (Steenstrup MS), Hoyle (1886) ;
Octopus rugosus (Bosc), Robson (1929). Challenger Coll., Ascension, April 1878.
C. 69.
This specimen, an immature female, is the type of 0. occidentalis and was described
by Hoyle (1886) in the Challenger Reports. In its present state it is very dark and
hard. A comparison of its present measurements with those given by Hoyle (Table
I) indicate that a very considerable shrinkage of the arms has taken place during
sixty years of preservation (the specimen was examined by me in 1946) but the
length and proportions of the mantle and head are little changed. The longest arm,
the second left, now measures about 180 mm. whereas Hoyle gives a length of
230 mm. No doubt his measurements, which represent the condition of the specimen
when it was in a better state of preservation, should be accepted.
Robson could find no reason why this specimen should not be assigned to 0.
rugosus and I can see no reason why it should not be assigned to 0. vulgaris.
West Africa
(9) B.M., 1928.3. 17. 1. Octopus rugosus (Bosc), Robson (1929). Old Collection
(Rich), West Africa. Bears manuscript name " Octopus rugosus var. longi-
manus. Type." C. 260.
This is a mature male, very dark in color. The bodily proportions are those of
vulgaris (Table I) the ligula (Table II) is minute and typical of vulgaris. The gills
are very damaged and the low estimate of the number of gill lamellae may be due
to this condition. Spermatophores were removed from Needham's organ ; the
length (9 mm.) gives an index of 37-5, which is typical of vulgaris. The horn is
very slender and measures about 4 mm. in length ; there are 40 to 50 spiral turns,
partly uncoiled ; only the distal extremity of the horn, proximal to the middle
piece, is not coiled. This last mentioned feature is not characteristic of a well pre-
served spermatophore of vulgaris but the partial uncoiling and poor state of preserva-
tion may well account for such a condition (compare Adam, 1952, Fig. 53B). There
seems to be no reason why this specimen should not be assigned to 0. vulgaris.
(10) B.M., 1928.3. 17.2. Octopus rugosus (Bosc), Robson (1929). Sierra Leone,
W. P. Lowe. Bears manuscript name " Octopus rugosus ? var. longimanus."
C. 262.
A juvenile specimen, rather warty, with 4 dorsal papillae in a rhomb on the back.
Probably referable to 0. vulgaris.
(11) B.M., 1928. 3. 17. 3. Octopus rugosus (Bosc), Robson (1929). Murray Town,
Sierra Leone, W. P. Lowe. C. 371.
REVISION OF OCTOPODINAE IN THE BRITISH MUSEUM 157
This juvenile specimen is very dark and extremely hard and shrunken. It is
very probably a young vulgaris.
(12) B.M., 1889.4.24. 15. Octopus tuberculatus Blainville, Hoyle (1886) ; Octopus
rugosus (Bosc.), Robson (1929). Challenger Coll., No. 122B, off Barra Grande,
Lat 9 9' S., Long. 34 53' W., depth 32 fm, red mud, 10.ix.1873. C. 115a.
This specimen was identified as 0. tuberculatus by Hoyle, but is not described.
There are indications of the dorsal rhomb of papillae and the specimen is very
probably referable to 0. vulgaris.
South Africa
(13) B.M., 1890.9.4.4. Octopus rugosus (Bosc), Robson (1929). Port Elizabeth,
South Africa, H. S. Spencer. Bears manuscript label " Octopus granulatus
Lam." C. 108.
A mature male with all the characteristics of 0. vulgaris. The specimen is very
hard but one spermatophore was recovered from Needham's organ and was found
to be in a moderately good state of preservation. Its length (10-5 mm) gives an
index that is typical of vulgaris (Table II). There are about 37 coils to the spiral
Table II. — Characteristics of adult male specimens referable to Octopus vulgaris Lam*
B.M. No.
West Agrica
1928.3. 17. 1
South Africa
1890.9.4.4 .
1924.9.9. 11
* Symbols as in Pickford (1945).
horn, interrupted by two irregular regions of partial uncoiling. The direction of the
spiral is reversed several times, as follows : 8 coils anticlockwise, about 6 irregular
clockwise coils, an irregular break, 3 anticlockwise coils followed immediately by 5
clockwise turns, an irregular break, 8 anticlockwise coils followed by 6 clockwise.
This repeated reversal of the spiral is not a characteristic feature of vulgaris spermato-
phores but there has been no large scale survey of spermatophore variation in this
species and, in the absence of such data and of an adequate series of spermatophores
from South African males, it seems unwise to attach any great significance to the
condition. Apart from this feature the specimen appears to be a typical 0. vulgaris.
(14) B.M., 1924.9.9. 11-13. Octopus rugosus (Bosc), Robson (1924 and 1929).
Natal, Gilchrist Stns. 388 and 389. C. 105-107.
These specimens were identified but not described by Robson. The male has a
minute ligula (Table II) and is apparently a typical vulgaris ; there is one broken
spermatophore in Needham's organ but it was not removed. The two females have
ripe ovaries with small eggs. The specimens are rather small in size for sexually
LLI
CLI
PLI
SpLI
Horn Coils
i-8
28
. i8- 7
37'5
. 40-50
0-9
50
16-7
35
. ca.sj
i-8
30
—
—
—
158 REVISION OF OCTOPODINAE IN THE BRITISH MUSEUM
mature females of vulgaris but are apparently referable to this species and resemble
the male with which they were associated.
(15) B.M., 1926.10.20.30. Polypus granulosus (Lamarck), Massy (1927) ; Octopus
rugosus (Bosc), Robson (1929). Cape Colony, South African Museum 1311D.
C. 109.
This female was described by Massy and indices based on her measurements are
given in Table I. The specimen closely resembles the South African specimens from
Natal, listed above, and similarly can be referred, in all probability, to 0. vulgaris.
(16) B.M., 1889.4.24. 18. Octopus granulatus Lam., Hoyle (1886) ; Octopus rugosus
(Bosc), Robson (1929). Challenger Coll., Simon's Bay, Cape of Good Hope,
10-20 fms. C. 114.
A very young specimen, with dorsal papillae in a characteristic rhomb, very
probably referable to 0. vulgaris.
In addition to the specimens listed above Robson's type of Octopus rugosus var.
sanctae helenae (B.M., 1868. 3. 12. 1) was examined but nothing new can be added
to the original description (Robson, 1929) . Although it is a badly distorted specimen
with somewhat deceptive features there seems to be no valid reason why it should
not be assigned to 0. vulgaris. The same is true also of Octopus verrucosus Hoyle
(1886) (B.M., 1889.4.24. 16). The type of this species was fully redescribed by
Robson (1929) who states that " It seems to exhibit characters of 0. vulgaris and
0. rugosus in combination with individual peculiarities." The distinctive features
listed by Robson do not appear to be sufficiently diagnostic to justify recognition
of this species as distinct from 0. vulgaris.
The following summary is offered to clarify the present synonymy of 0. vulgaris,
in the light of the considerations outlined above.
Octopus vulgaris Lamarck
Octopus vulgaris, Robson (1929). All specimens from the north-eastern, north-
western, central and south Atlantic, and from West and South Africa, which
Robson refers to this species must be accepted as correctly identified. Speci-
mens from the Indo-Pacific require further confirmation.
Octopus rugosus, Robson (1929). The majority of specimens from the eastern,
western, central and south Atlantic, and from West and South Africa which
Robson referred to this species are probably correctly assigned to 0. vulgaris.
It cannot be established that 0. rugosus is a valid species. Some American
specimens are referable to 0. briar eus Robson, and 0. joubini Robson (Pickford,
1945, 1946), and one specimen from the Cape Verde Islands, described as a new
species in the present contribution, must be excluded.
Octopus rugosus var. sanctae helenae, Robson (1929) = 0. vulgaris.
Octopus verrucosus, Hoyle (1886), Robson (1929) = 0. vulgaris.
Octopus vulgaris, Pickford (1945, 1946). The proposed synonymies of western
Atlantic species appear to remain valid. 0. geryonea Gray, 0. eudora Gray,
REVISION OF OCTOPODINAE IN THE BRITISH MUSEUM 159
0. filosus Howell, 0. carolinensis Verrill, and various other obscure western
forms are assimilated into the synonymy of 0. vulgaris.
Octopus vulgaris, Adam (1952). Probably all the West African specimens which
Adam referred to this species are correctly assigned. No doubt would arise if
it were not for the two specimens listed from the Cape Verde Islands, but not
identified, which must be placed in a new species (see below).
There is an urgent need for the re-examination of specimens assigned either to 0.
vulgaris or to 0. rugosus, granulatus or tuberculatus from the Indo-Pacific. As Robson
has pointed out, the ligula of oriental specimens of 0. rugosus (as understood by him)
differs from that of Atlantic specimens ; the average size is larger and the calamus
shorter. A consideration of the data, so carefully assembled by Robson, appears to
indicate that the Indo-Pacific rugosus-complex is composite in character and
distinct from the Atlantic rugosus-complex. It is difficult to understand why he
discarded this interpretation but one factor was the confusion that arose due to the
inclusion of western Atlantic species that are not 0. vulgaris in the synonymy of
0. rugosus. The range of variation in critical characters, such as the length of the
ligula, was thus enlarged in such a manner as to obscure the true situation.
Octopus vincenti n. sp.
Octopus granulatus (part), Hoyle (1886).
Octopus rugosus (part), Robson (1929).
Octopus sp., Adam (1952).
Holotype : lies du Cap- Vert, 28.x. 1948, Exped. Oceanogr. Beige cotes africaines de l'Atlantique
Sud. Inst. Royal Sci. Naturelles de Belgique.
Specimen examined :
B.M., 1889.4. 24. 17. Octopus granulatus Lam., Hoyle (1886) ; Octopus
rugosus (Bosc), Robson (1929). Challenger Coll., St. Vincent, Cape Verde
Islands, 25. iv. 1876, depth 15-20 fms. C. 118.
In 1952 Adam listed a male and female from the Cape Verde Islands which were
obviously different from Octopus vulgaris, but to which he hesitated to give a new
name. The length of the ligula in the male is quite outside the range of vulgaris
and the horn of the spermatophore is straight, without indications of uncoiling
(Adam, Fig. 53D). These two features alone are sufficient to justify the recognition
of a new species since no such males have hitherto been described from the eastern
Atlantic. In addition the animal is of small size, mantle length 25-29 mm., the arms
are short, the web deep and the suckers small, close to the lower limits for 0. vulgaris.
The female has a similar combination of characters but, on account of the great
variability of Octopus vulgaris, Adam concluded that it was not possible to recognise
the female as a distinct species.
During the course of the present investigation it was discovered that the male
listed above, in the Collections of the British Museum, presented the same combina-
tion of characters. It was identified by Hoyle as 0. granulatus and referred by
Robson to 0. rugosus, but under no circumstances can it be placed, along with other
eastern Atlantic specimens of Robson's rugosus list, in the synonymy of 0. vulgaris.
160 REVISION OF OCTOPODINAE IN THE BRITISH MUSEUM
The size of the ligula alone precludes such an identification. The similarity to Adam's
male is so great that there can be no doubt that the two specimens are conspecific.
The only special feature of the British Museum specimen is the presence of a faintly
darker area in front of the eye, about 4 mm. in diameter, which could possibly be
interpreted as the remains of a faded ocellus ; but this interpretation seems unlikely.
The British Museum specimen is not sexually mature and, in view of the highly
distinctive spermatophores of Adam's specimen, which were in a good state of preser-
vation and have been carefully figured, it seems preferable to designate this male
as the holotype. The female, also from the Cape Verde Islands and with similar
bodily features, is almost certainly referable to the same species but caution dictates
that it should not be designated as the gynotype. It is unfortunate that the ovary
was not sufficiently advanced for the size of the eggs to be determined.
The bodily proportions and characteristics of the three specimens are summarized
in Table III. Adam did not describe either the funnel organ or the penis. Un-
fortunately the British Museum specimen does not allow one to give any account
Table III. — Octopus
vincent i n. sp.
Tab
ular comparison of specimens.
Specimen
ML MWI HWI
MAI
WDI SnI Gill LLI CLI SpLI
Holotype : $ (Adam, 1952)
$ (Adam, 1952)
BM.1889.4.24.17, 6"
29 55 59
25 72 60
25 88 68
50*
40*
42
34 9-5 10 6-2 40 55f
22 9-0 10 — — —
28 8-o 9 5 -6 40 —
* Calculated from Adam's index (Longest arm /Mantle length).
f Calculated from Adam's Fig. 53 D x + ^V It is not quite clear whether the two sections, horn and
sperm reservoir respectively, show the complete spermatophore or whether a middle piece has been
omitted. If this is the case the spermatophore length index would be greater.
of these characters. The funnel organ is too poorly preserved for description. The
specimen had already been dissected and no sign of the penis remains ; it was
probably small and undeveloped since the animal is not sexually mature. There are
no spermatophores.
Diagnosis : A medium-sized species (mantle-length 24-29 mm.) with granular
skin and bodily proportions similar to those of immature specimens of 0. vulgaris.
The arms are short, 2 to 2-5 times the length of the mantle, the first arms shortest.
The web is deep, about one-third the length of the longest arm, the D sector deepest,
the A sector shallowest. The suckers do not exceed 10% of the mantle-length and
there are no specially enlarged suckers in either sex. The funnel organ is not known.
The gills have 9 or 10 primary lamellae in each demibranch ; the high number
resembling that found in 0. vulgaris. In the male the third right arm is hectocoty-
lized and the ligula represents 4-5-6-2% of its length, the calamus opens near the
middle, two-fifths of the distance from the last sucker to the tip of the ligula. The
spermatophore is about half the length of the mantle and the horn is straight,
without coils. The size of the eggs is unknown.
It may be enquired whether any other specimens, either among those listed by
Adam or in the series from the British Museum can reasonably be assigned to this
species. British Museum males have the characteristic minute ligula of 0. vulgaris.
REVISION OF OCTOPODINAE IN THE BRITISH MUSEUM 161
A juvenile male from the Cape Verde Islands (B.M., 1855.9.29.2) can also be
excluded. The arms are longer and the web less deep, moreover the state of im-
maturity at a mantle-length of 31 mm. indicates that it is the young of a large
species. The immature female from the Cape Verde Islands (B.M., 1851 . 1 . 24 . 1) with
a mantle-length of 50 mm. may be excluded for similar reasons, including also the
larger relative size of the suckers. Among Adam's specimens there are two males
that have an unusually large ligula for 0. vulgaris (Goree, 10. vi. 1947, LLI 3*5 ;
Goree, 24. ix. 1946, LLI 2-9) but the spermatophore of the latter, illustrated by
Adam (Fig. 53A), is typical of 0. vulgaris, with a spirally coiled horn. Moreover in
both these specimens the arms are long, the web only of moderate depth, and the
suckers large as in 0. vulgaris. Among females there is one (Goree, 31^.1950) with
unusually short arms, a moderately deep web, and small suckers, that might well
be referred to this species. However, the characters of the females present no clear
cut distinctions by which one species could be separated from the other. A clarifica-
tion of the problem must await the capture of a gravid specimen ; it may be antici-
pated that the ripe eggs may show some distinctive features.
It is unlikely that the sexually mature male of 0. vincenti could be confused with
any other Atlantic species. The differences between this species and 0. vulgaris
have already been discussed. In the western Atlantic there are four Caribbean
species that have a ligula of approximately the same proportions as in 0. vincenti.
0. briareus Robson is readily excluded, the arms are long, the number of gill lamellae
low, and the spermatophore horn is coiled. 0. joubini Robson is a nearly smooth-
skinned species with short arms, the spermatophore horn is straight but the number
of gill lamellae is low, well below the minimum recorded for 0. vincenti. The ocellate
species, 0. hummelincki Adam, has, in addition to this distinctive feature, a low
number of gill lamellae and a coiled spermatophore horn. The nearest affiliation
would appear to be with 0. burryi (Voss, 1950, 1951). This species has a characteristic
color pattern in the form of a purplish band which extends the full length of each
arm on the inner (dorsal) side. Apart from this highly distinctive feature, it shows
a general resemblance to 0. vincenti and, in preparing a key for the separation of
the two species it would be necessary to depend primarily upon the presence or
absence of the purple arm band. Thus the arms are short, the web deep, and the
number of gill lamellae high. The ligula is similar in size to that of 0. vincenti and
the horn of the spermatophore is nearly straight with only a few spiral turns (one
proximal and three distal in the one that has been described).
The problem of 0. rugosus in the Indian Ocean is so little understood that it would
be futile, at the present time, to offer any comparison with these or other oriental
forms.
My best thanks are due to Dr. W. J. Rees for facilities to study the British
Museum specimens.
SUMMARY
i. Twenty- two specimens from the eastern Atlantic, the west African coast and
South Africa which Robson (1929) identified as 0. rugosus and which are now in
the Collections of the British Museum have been re-examined.
162 REVISION OF OCTOPODINAE IN THE BRITISH MUSEUM
2. Fifteen are either juvenile or sexually immature but, with one doubtful excep-
tion, they can reasonably be assigned to 0. vulgaris Lam.
3. One juvenile specimen from Madeira has a peculiar zebra pattern on the arms
and may therefore represent the young of some different species.
4. Six of the 7 adults show no features which would justify their separation from
0. vulgaris Lam.
5. One adult male from the Cape Verde Islands is assigned to a new species. A
male and female of this species were described but not named by Adam (1952) and
Adam's male has been designated as the Holotype of Octopus vincenti n. sp.
6. The nature of the South African vulgaris-rugosus population is discussed and it
is concluded that, at most, the South African specimens may represent a local race
of slightly smaller size and with other minor differences, not specifically separable
from 0. vulgaris Lam.
REFERENCES
Adam, W. 1952. C6phalopodes. Exp6d. Oceanogr. Beige dans les Faux C6tieres Africaines
de l'Atlantique Sud, Res. Sci. 3 (3) : 1-142, pis. 1-3.
Bosc. L. 1792. Sepia rugosa. Actes Soc. d'Hist. Nat. Paris, 1 (1) : 24, PI. V, figs. 1-2.
Hoyle, W. E. 1886. Report on the Cephalopoda collected by H.M.S. " Challenger " during
the years 1873-76. Rpt. Sci. Res. Voyage, H.M.S. " Challenger," Zool. 16 : 1-246, pis. 1-33.
Lamarck, J. B. 1799. Sur les genres de la Seche, du calmar et du poulpe. Mem. Soc. Hist.
Nat. Paris, 1 : 1-25.
Massy, A. L. 1925. On the Cephalopoda of the Natal Museum. Ann. Natal Mus. 5 : 201-230.
1927. The Cephalopoda of the South African Museum. Ann. S. A jr. Mus. 25 : 1 51-168.
Pickford, G. E. 1945. Le poulpe am6ricain : A study of the littoral Octopoda of the western
Atlantic. Trans. Conn. Acad. Arts Sci. 36 : 701-81 1, pis. 1-14.
1946. A review of the littoral Octopoda from central and western Atlantic stations in the
collections of the British Museum. Ann. Mag. Nat. Hist. (11) 13 : 412-429.
Robson, G. C. 1924. On the Cephalopoda obtained in South African Waters by Dr. J. D. F.
Gilchrist in 1920-21. Proc. zool. Soc. London, 1924 : 589-686.
1929. A Monograph of the Recent Cephalopoda. Part I — Octopodinae. 236 pp., pis. 1-7.
London (Brit. Mus.).
Voss, G. L. 1950. Two new species of cephalopods from the Florida Keys. Rev. Soc. Mala-
cologica 7 (2) : 73-79, figs. 1-3.
1951. Further description of Octopus burryi Voss with a note on its distribution. Bull.
Marine Sci. Gulf Carib. 1 (3) : 231-240, figs. 1-6.
REVISION OF OCTOPODINAE IN THE BRITISH MUSEUM 163
Part II.— A RE-EXAMINATION OF THE SPECIMENS OF
ROBSONELLA FONTANIANA IN THE COLLECTIONS
OF THE BRITISH MUSEUM
Robsonella (= Joubinia) fontaniana (Orbigny) is a well defined species, endemic
to the subantarctic or antiboreal region of South America. Supposed records of this
species from the Indian Ocean and from the Central Pacific are subject to the gravest
suspicion, as pointed out by Robson (1929). Massy (1925) described a specimen from
Natal which Robson has considered to be a local variety. Two closely related but
supposedly distinct species have been described from Australia and New Zealand,
viz. R. australis (Hoyle) and R. huttoni Benham. Polypus campbelli of Smith is
included in the synonymy of R. australis (Robson, 1929, footnote p. 145).
Robson (1929) placed fontaniana and australis (+ campbelli) in the new genus
Joubinia but this was shown to be pre-occupied and the name Robsonella was sub-
stituted by Adam (1938). The characters of the genus were summarized by Robson
and have been reviewed recently by Dell (1952). They are as follows : the arms are
subequal, the web is deep and subequal, the mantle aperture is partly closed, the
adlateral tooth of the radula is supposedly bicuspid, the penis has a long primary
and a small secondary diverticulum, and the ligula of the hectocotylus is remarkably
stout with inrolled sides and wide cheeks. We may disregard the characters of the
arms and web since these are not of generic value. The partial closure of the mantle
is also unimportant since many species of the genus Octopus have a slightly narrowed
mantle opening of Robson's type B, moreover the mantle aperture is wide in R.
australis (Benham, 1942). The long diverticulum of the penis is highly distinctive as
a species character, common to all three forms, but there are species of Octopus which
show a similar feature. The secondary diverticulum to which Robson attached great
significance is apparently a variable feature since it was lacking in one of Dell's
specimens of R. australis. There remains only the remarkably stout form of the ligula
which may be conveniently retained, at least for the present, as a generic character.
However it appears probable that the genus Robsonella is not a valid one and that
the species which have been assigned to it should be returned to the genus Octopus.
A second problem concerns the validity of the distinction between R. fontaniana
and the two Australasian species. Time did not permit a re-examination of the
specimens of R. australis (-{- campbelli) which are in the collections of the British
Museum, but the material has been well described and we have also the accounts
of Benham (1942) and Dell (1952) for comparison. The entire series of n specimens
of R. fontaniana was carefully re-examined with a view to confirming and enlarging
our concept of this South American species (Tables I and II). There are insignificant
differences between the mean values of the indices as given by Robson (1929) and
as determined here. In addition there are some features which call for further
discussion.
164
REVISION OF OCTOPODINAE IN THE BRITISH MUSEUM
Table I. — Robsonella fontaniana (Orbigny). Bodily characteristics of eleven specimens
in the Collections of the British Museum .*
B.M. No.
Sex
ML
MWI
HWI
MAI
WDI
SnI
Sel
Gill
1851.1.24.5 .
0"
32
84
75
27
19
11 -o
17-2
9-5
1869.6.5.54 .
6*
24
108
7i
34
32
IO-2
14-6
9
1869.6.5.65 .
6*juv.
19
84
68
28
16
10-5
13-2
ca. 8
1869.6.5.69 .
6* juv.
19
100
55
35
18
6-8
9-5
10
1851.1.24.12
?mat.
24
92
67
39
32
12-5
—
10
1848.6. 16.2 .
?
35
69
54
32
21
IO-O
—
9
1869.6.5.63 .
?
27
78
63
34
20
7*4
—
9'5
1868.7. 10.2 .
?
25
72
68
42
32
11-4
—
10-5
1899. 8. 31 .84
?
22
96
87
39
25
9-1
—
9
1869.6.5.62 .
?
22
104
73
37
30
II-4
—
ca.g
1851.1.24.4 .
juv.
10
100
90
42
29
io-o
—
10
Average
99
77
37
25
11 -o
I3'8
9'5
* Symbols as in.
Pickford
(i945).
Table II.-
B.M. No.
1851.1.24.5
1869.6.5.54
1869.6.5.65
1869.6.5.69
-Robsonella fontaniana (Orbigny). Characteristics of male specimens in
the Collections of the British Museum*
SpLI Sp. Horn Comments
165 No coils —
83 One turn —
(not formed) Immature
(not formed) Immature
LLI CLI
7-8 50
5-i 50
(damaged)
3*9 56
PLI
23-5
27
18-5
15-8
* Symbols as in Pickford (1945).
Robson's sucker-diameter index does not distinguish between the diameter of the
largest suckers of the normal series and the diameter of the specially enlarged
suckers. It was found that such specially enlarged suckers occur only in males and
are present in the four representatives of this sex that were examined. The penis of
the two adult males is as figured by Robson, with an accessory knob-like diverti-
culum. The penes in the two immature specimens are similar but they lack the
secondary diverticula. The ligula-length index averages 6 • 5 in the two adult speci-
mens ; as might be expected it is somewhat smaller in an immature male.
Two spermatophores were found in the larger male (B.M., 1851.1.24.5), lodged in
Needham's organ, and one was removed for study. It is poorly preserved and
doubled upon itself in a knot in the middle region. The estimated length of the
horn plus middle piece is 38 mm., the sperm reservoir measures 15 mm. The total
length is thus half as long again as the mantle (SpLI 165). The horn is dark and
opaque and there is no evidence of coiling. Towards its distal end, i.e. proximal to
the knot-like tangle, the inner tube has a moniliform appearance due to a series of
three constrictions which are probably artifacts of preservation.
A single spermatophore was found in the smaller male (B.M., 1869.6.5.54). The
horn plus middle piece measure 13 mm. and the sperm reservoir measures 7 mm.
REVISION OF OCTOPODINAE IN THE BRITISH MUSEUM 165
The spermatophore-length index is thus lower than in the larger specimen (SpLI 83) .
The horn is straight except for a single widely-open turn of a spiral towards its
distal end.
Nearly all the females are sexually immature with small and relatively undeveloped
ovaries. However, the reproductive ducts are well developed and have been des-
cribed by Robson. It may therefore be inferred that the specimens are spawned
out or in a state of sexual quiescence rather than immature. Robson calls attention
to the distal portion of the oviduct, or vagina, which is long and stout, but I find
that this feature is somewhat variable. One specimen (B.M., 1851.1.24.12) has a
well developed ovary with moderately large-sized eggs, ca. 3*5 mm. in length. The
eggs are not fully ripe and the length of the stalk could not be determined. The
vagina of this specimen is not abnormally stout.
DISCUSSION
Table III gives a comparison of indices and numerical characteristics of the
species of Robsonella, compiled from data presented by Robson (1929), Benham
(1942, 1943), and Dell (1952), and from the re-examination of the British Museum
series of R. fontaniana. This table demonstrates the difficulty of assessing the
Table III. — Comparison of the species that have been assigned to the genus Robsonella.*
ML
MWI
HWI
ALI
WDI
SnI
Sel+
Gill
LLI
Species
range
av.
av.
av.
av.
av.
av.
range
range
R . fontaniana (Orbigny)
B.M. Seriesf .
10-35
99
77
(74)
25
11 -o
i3'8
8-10
5-i-7-8||
R. australis (Hoyle)
Robson (1929)
australis
22-33
81-86
62-73
72-76
33
11-13
—
6-9
io-8
" campbelli "
28
85
7i
78
28
?
21
10
8-5
Benham (1942)
10-36
79
62
73
24
12
—
6-8
8-4
Dell (1952)
11-30
76
63
73
27
io*6
—
6-8
4-5-4-7
R. huttoni (Benham)
Dell (1952)
43-56
68
44*5
74
22
9'4
—
6-7
6-6-6-8
* Symbols as in Pickford (1945). Doubtful specimens from the Indian Ocean and Central Pacific
excluded.
f Based on new measurements except for the arm-length index which is taken from Robson (1929)
and which had to be used, instead of the mantle-arm index, to permit comparison with the published
data for the other forms.
I Specially enlarged suckers found only in males.
|| Sexually mature specimens only.
differences between the three species. R. huttoni differs from R. australis in its
larger size, narrower body and much narrower head but a familiarity with the
complexities of octopodan taxonomy suggests that these features are difficult to
evaluate. The same is true of the relatively somewhat smaller size of the suckers.
Both these Australasian species differ from R. fontaniana primarily in the smaller
number of gill lamellae ; the ranges, however, overlap and the type of " campbelli "
166 REVISION OF OCTOPODINAE IN THE BRITISH MUSEUM
has fully as many lamellae as a typical specimen of R. fontaniana. It must be
remembered that an accurate count of the number of primary lamellae is highly
subjective, depending upon the number of minute terminal foliations that are
included in the count. However, the difference appears to be a valid one and must
be accepted as such in the present state of our knowledge of these species. As
indicated by Dell (1952) R. huttoni will probably prove to fall within the range of
R. australis, but it is also evident that the Australasian specimens are at least
racially distinct from the South American form.
A few more points may be discussed. The eggs of R. australis measure 2 • 5 mm.
in length and have a short stalk (Benham, 1942). The eggs of R. huttoni are of about
the same size, 3 mm. (Benham, 1943). Eggs, similar in size, measuring about 3 • 5 mm.,
are present in the ovary of a mature female of R. fontaniana, described here.
The characters of the hectocotylus and penis have been fully discussed by previous
investigators and appear to offer no evidence for separating the species. Attention
may be called to a curious error in Dell's paper, in discussing the affinities of R.
huttoni he states on p. 40 that " The ligula index is appreciably lower than in
australis — average 6-7 as compared with 10 • 8." But his table (p. 34, Table 9) shows
that his own specimens of R. australis have a low index, averaging 5*4. The specimens
appear to have been sexually mature since the presence of a spermatophore is
mentioned in at least one of them.
The spermatophores of R. fontaniana are described for the first time. They are
nearly as long as the mantle, or considerably longer. The horn is straight, as far as
could be determined, or with a single distal turn, but admittedly the preservation
is not good. Benham described and figured the spermatophores of R. australis. He
states that they measured 40 mm. in length. The mantle-length of the male from
which these spermatophores were taken is not given, but even his largest specimen
has a mantle-length of only 36 mm. Therefore the spermatophore, like that of R.
fontaniana, is about as long as or longer than the mantle. In the text Benham states
that the " projectile apparatus " consists of a " closely-coiled spring". This would
lead one to suppose that the horn of the spermatophore was coiled. However, a
study of his figures (Benham, 1942, Figs. 6 and 7) suggests quite otherwise. The
horn is apparently straight and what Benham mistook for the coils is clearly, in
reality, the internal structure of the lumen which always has this delicate spiral
structure, irrespective of whether the horn is straight or coiled (compare with the
spermatophores of 0. macropus, figured by Pickford, 1945, PI. IV). The spermato-
phores of R. huttoni have not been described.
The author is indebted to Dr. W. J. Rees, for facilities to study these specimens.-
SUMMARY
i. Eleven specimens of Robsonella fontaniana (Orbigny) in the Collections of the
British Museum have been redescribed. The spermatophores, described for the first
time are nearly as long as or longer than the mantle ; the horn is straight or with a
single spiral turn towards its distal end. The eggs are moderately large, ca. 3*5 mm.
in length.
REVISION OF OCTOPODINAE IN THE BRITISH MUSEUM 167
2. The status of the genus Robsonella is discussed. It is considered probable that
the two Australasian species, R. australis (Hoyle, 1885) and R. huttoni (Benham,
1943) are synonymous but together they form an assemblage that is at least racially
distinct from the South American species, R. fontaniana. The chief distinguishing
characteristic is the lower average number of primary gill lamellae in the Australasian
species.
REFERENCES
Adam, W. 1938. Robsonella nom. nov. fur Joubinia Robson, 1929. (Cephalopoda, Octopoda).
Zool. Anz. 121 : 223.
Benham, W. B. 1942. The octopodous Mollusca of New Zealand : (1) The midget Octopus
of coastal waters. Trans. R. Soc. New Zealand, 72 : 226-236.
1943- The octopodous Mollusca of New Zealand. — Part II. Trans. R. Soc. New Zealand,
73 : 53-57.
Dell, R. K. 1952. The Recent Cephalopoda of New Zealand. Dominion Mus. Bull., No.
16 : 1-157.
Massy, A. L. 1925. On the Cephalopoda of the Natal Museum. Ann. Natal Mus. 5 : 201-230.
Pickford, G. E. 1945. Le poulpe americain : A study of the littoral Octopoda of the western
Atlantic. Trans. Conn. Acad. Arts Set. 36:701-811.
Robson, G. C. 1929. A Monograph of the Recent Cephalopoda. Part I — Octopodinae. London
(Brit. Mus.).
PRINTED IN GREAT BRITAIN BY
AD LARD AND SON, LIMITED,
BARTHOLOMEW PRESS, DORKING
2 SEP 1955
A REVISION OF THE FAMILY
EPICRIIDAE
(ACARINA— MESOSTIGMATA)
G. OWEN EVANS
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 3 No. 4
LONDON: 1955
A REVISION OF THE FAMILY EPICRIIDAE
(ACARINA— MESOSTIGMATA)
BY
G. OWEN EVANS
Pp. 169-200 ; Plates 1-2 ; 41 Text-figures
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 3 No. 4
LONDON: 1955
THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, is
issued in five series corresponding to the Departments
of the Museum, and an Historical Series.
Parts will appear at irregular intervals as they become
ready. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.
This paper is Vol. 3, No. 4 of the Zoological series.
PRINTED BY ORDER OF THE TRUSTEES OF
THE BRITISH MUSEUM
Issued August, 1955 Price Ten Shillings
A REVISION OF THE FAMILY EPICRIIDAE
(ACARINA— MESOSTIGMATA)
By G. OWEN EVANS
CONTENTS
Page
Introduction ........... 171
Classification . . . . . . . . . . -173
Genus Epicrius Can. & Fanz. . . . . . . . -174
Epicrius mollis (Kramer) . . . . . . . 175
Epicrius canestrinii Haller . . . . . . . .183
Epicrius menzeli Schweizer . . . . . . . .185
Epicrius minor Willmann . . . . . . . .188
Epicrius spinituberculatus n.sp. ....... 188
Species Dubiae . . . . . . . . . .191
Genus Berlesiana Turk . . . . . . . . . .192
Berlesiana cirratus (Berl.) . . . . . . . -193
Berlesiana denticulata n.sp. ........ 193
Geographical distribution of the Epicriidae ..... 196
Discussion ........... 196
Summary ............ 198
Acknowledgments .......... 198
References ........... 198
INTRODUCTION
The family Epicriidae Berl. comprises a small group of Mesostigmatid mites readily
distinguished in the adult stage by the polygonal network of bi- or tri-iurcate tuber-
cles on the dorsal shield and the absence of an ambulacral apparatus on the first
pair of ambulatory appendages. This characteristic ornamentation of the dorsum
(" seriebus papillarum fuscarum symmetrice reticulato ") was first described as
occurring in Gamasus reticulatus Grube, 1859, but unfortunately the original descrip-
tion is inadequate for the recognition of its specific identity. In 1877 Canestrini
& Fanzago figured a species for which they proposed the name Epicrius geometricus,
thereby also erecting the new genus Epicrius. Later, Canestrini (1885) added a
short description of the species. In the meantime, Haller (1881) had given a defini-
tion of Epicrius and added two other species to it, Gamasus mollis Kramer, 1876 and
Epicrius canestrinii Haller, 188 1. His description and figures of the latter are
sufficiently detailed for recognition of the species.
Berlese (1886-87) in "Acari, Myriopoda et Scorpiones hucusque in Italia reperta "
re-described and figured E. geometricus stating that it was found " in totius Italiae
muscis, praecipue in montibus." In the same work E. canestrinii is referred to as a
ZOOL. 3, 4. 14
172 A REVISION OP THE FAMILY EPICRIIDAE
variety of geometricus. Four other species, Epicrius corniger BerL, Epicrius glaber
BerL, Epicrius laelaptoides BerL and Epicrius mollis (Kramer) Berl. were also con-
sidered to be congeneric with E. geometricus. Later, Berlese (1916a) emended his
concept of the genus Epicrius and transferred corniger, glaber and laelaptoides to the
genus Lasioseius s. lat., and mollis to the genus Epicriopsis Berl. Further, in Berlese
(1916&) Gamasus mollis Kramer 1876 non Berl., 1887 was considered to be the nymphal
stage of E. geometricus which therefore falls into synonymy. Three new species of
Epicrius were also described in this paper, namely, E. cirratus, E. washingtonianus
and E. parisiensis. The latter was made the type of a new subgenus Di epicrius.
Absolon (1899) erected the genus Eugamasus (preoccupied by Eugamasus BerL,
1893) for Eugamasus cavernarum Absolon found on the excretory material of bats in
a cave in Moravia. There is no doubt that this species is an Epicriid, probably of the
genus Epicrius. It does not appear to have been found subsequently.
George (1906) re-described three species of Epicrius from Lincolnshire. The
species he identified as Epicrius mollis (Kramer) is Epicriopsis horridus (Kramer)
and what he believed to be Epicrius canestrinii has recently been given the new name
Cornubia georgei by Turk (1943) . The specimens of the third species are in the collec-
tions of the British Museum (Natural History) and are correctly determined as
Epicrius geometricus Can. & Fanz. (= E. mollis).
Schweizer (1922) listed two species of the genus Epicrius from Switzerland,
E. geometricus and Epicrius menzeli Schweizer, 1922. The specimens of the former
were collected from three main regions, namely, " Mittelland," " Jura " and" Alpen".
The females described from " Mittelland " and " Alpen " were correctly determined,
but the female described and figured from the " Jura " was, as will be shown below,
Epicrius canestrinii Haller.
Tragardh (1942) in his revision of the genus Epicrius unfortunately deals only
with E. mollis. This paper is important for the historical account of the genus and
the description of the larva and the protonymph of the genotype. In addition,
Tragardh has dealt with the classification of the family Epicriidae in several papers
on the comparative morphology of the Mesostigmata e.g. Tragardh 1938, 1946a & b.
Turk (1943) has also added to our knowledge of the Epicriidae in his work on the
British species. He erected a new genus, Berlesiana, for Epicrius cirratus Berl. on
the basis of the structure of the sternal region in the female. The genus Cornubia,
which was also proposed in this paper, was later made a synonym of Ameroseius BerL
(Turk, 1953). Two species are included in the genus Epicrius, namely, E. mollis
and E. geometricus. His E. mollis, however, is Epicriopsis horridus (Kramer) and
this invalidates the discussion in that paper on the synonymy of E. mollis and E.
geometricus.
The most recent work on the genus Epicrius is by Willmann (1953). A new
sub-genus, Epicriella, was proposed for Epicrius [Epicriella) minor Willm.
The object of the present work is to review our knowledge of the classification of
the Epicriidae and to re-describe and figure, when possible, the known species.
This revision is based on material in the collections of the British Museum (Natural
History), the Oudemans Collection at Leiden and Dr. Jos. Schweizer 's Collection at
A REVISION OF THE FAMILY EPICRIIDAE 173
Basle. Dr. G. Lombardini has also kindly undertaken comparisons of specimens of
two species with the types in the Berlese Collection at Florence.
CLASSIFICATION
The supra-generic classification used in this paper follows that given by Evans
(1955) in which the Epicriina was divided into two super-families, the Epicrioidea
and Zerconoidea.
MESOSTIGMATA— EPICRIINA
" Mites with the epigynal portion of the genital plate in the female reduced to a
narrow chitinized rim overlapping the genital orifice. Male genital opening situated
in the sternal shield in the region of coxae II and III and closed by two plates, the
anterior of which bears a pair of hairs. Chelicerae dentate in both sexes but without
spermatophoral process in the male. Pedipalps with five free segments, specialized
seta on palptarsus two or three pronged" (Evans, 1955).
Super-family Epicrioidea Evans
Epicrioidea Evans, G. O. Bull. Brit. Mus. (nat. Hist.) 1954, Zool. 2 : 295.
Dorsal shield in both sexes provided with bi- or trifurcate tubercles forming a
polygonal network, and a pair of large dorso-lateral protuberances of unknown
function. Dorsal setae simple, setose or strongly barbed. Both sexes with jugularia;
sternal setae II and /or IV may be situated on a sclerotised shield or on the inter-
scutal membrane. Geniti-ventral shield in the female large, flask-shaped or rect-
angular. Sterniti-genital shield in the male extending posterior to coxae IV.
Gnathosoma with four pairs of ventral setae ; corniculi short, digitiform. Pedipalps
with the tibia and tarsus fused dorsally. Specialized seta on palp tarsus three-
pronged. Peritreme markedly reduced or absent. Stigmata enclosed in the lateral
extension of the dorsal shield. Leg I without ambulacral apparatus. Tarsus I (and
sometimes tibia I) with specialized, clubbed sensory setae. Legs II-IV with multi-
lobed pulvillus and two claws.
Family Epicriidae Berl.
Epicriidae Berlese, A. Bull. Soc. ent. Ital. 1885, 17 : 129.
The only family of the Epicroidea and therefore with the above characters. The
Epicriidae consists of two genera which may be distinguished as follows:
I. The majority of the dorsal setae more than ioo\l in length, simple or setose.
Dorso-lateral protuberances large, conspicuous. Tarsus I with three or
more clubbed sensory setae ...... Epicrius Berl.
-. The majority of the dorsal setae less than 60 \l in length, stout and strongly
barbed (fig. 32). Dorso-lateral protuberance small, inconspicuous.
Tarsus I with less than three pairs of clubbed sensory setae . Berlesiana Turk
i 7 4 A REVISION OF THE FAMILY EPICRIIDAE
I. Genus Epicrius Canestrini & Fanzago
Epicrius Canestrini, G. & Fanzago, F., Atti. 1st. Venet. 1877, (5) 4 pt. 1 : 131.
Eugamasus Absolon, Ph. C. K., Zool. Anz. 1899, 22 : 324.
Parasejus Tragardh, T. Naturw. Untersuch. des. Sarekgebirges. 1910, 4, 4 : 432.
Epicrius (Diepicrius) Berlese, A., Redia, 1916, 12 : 151.
Epicrius (Epicriella) Willmann, C, Sitzber. osterr. Akad. Wiss. math-nat. Kl. 1953, 1, 6 : 474.
Dorsal setae simple or setose ; the majority longer than 100/1. Dorso-lateral
protuberance large. Geniti-ventral shield flask-shaped. Tarsus I with three or
more clubbed setae. Other characters as in the definition of the super-family.
Type : Epicrius geometricus Can. & Fanz., 1877, (= Gamasus mollis Kramer, 1876).
This genus contains five species which may be considered valid and three species of
uncertain status pending the re-examination of the type material. The former may
be separated according to the following key to both sexes :
Key to the species of the genus Epicrius Can. & Fanz., 1877
Females.
1. Sternal setae II situated on the interscutal membrane between the jugularia and a
shield bearing sternal setae III and IV .... Epicrius minor Willmann
-. Sternal setae II situated on a well-sclerotised shield with setae III or setae III and
IV 2.
2. Dorsal setae D2 and D4 about one-third the length of setae D3 . . . . 3.
-. Dorsal setae D2 to D4 approximately equal in length. Sternal setae II to IV on an
undivided shield. Interscutal membrane between geniti-ventral and anal shields
with seven setae ........ Epicrius menzeli Schweizer
3. Projections of dorsal tubercles with rounded extremities (PI. 1) .... 4.
-. Projections of dorsal tubercles sharply pointed at their extremities (PI. 2). Sternal
setae II to IV on an undivided shield. Interscutal membrane between geniti-
ventral and anal shields with four setae. Anal shield with the usual three setae
Epicrius spinituberculatus sp. n.
4. The seta posterior to the large dorso-lateral protuberance separated from it by a
transverse row of tubercles. Sternal setae II to IV on an undivided shield. The
four setae between the genitiventral and anal shields each situated on a platelet 1
Epicrius canestrinii Haller
-. The seta posterior to the large dorso-lateral protuberance not separated from it by a
row of tubercles. Sternal setae IV (metasternals) separated from shield bearing
setae II and III. The pair of setae between the geniti-ventral and anal shields not
on platelets ........ Epicrius mollis (Kramer) Berl.
Males.
1. Anal shield completely fused with the dorsal shield. Posterior margin of the
sterniti-genital shield strongly convex. With three pairs of setae on the interscutal
membrane posterior to the sterniti-genital shield . . Epicrius mollis (Kramer) Berl.
-. Anal shield free, or partly fused with the dorsal shield. Posterior margin of the
sterniti-genital shield truncated . . . . . . . . . 2.
1 In some specimens the platelets on one side of the mid-ventral line may be fused so that there are
only three platelets between the geniti-ventral and anal shields, (see p. 13).
A REVISION OF THE FAMILY EPICRIIDAE 175
2. Setae D2 and D4 about one-third the length of setae D3 . . . . . 3.
-. Setae D2 to D4 approximately equal in length. Ventri-anal shield with ten pre-anal
setae .......... Epicrius menzeli Schweizer
3. Projections of dorsal tubercles rounded at their extremities. Region between the
sterniti-genital and anal shields occupied by a large rectangular shield bearing
three pairs of setae ....... Epicrius canestrinii Haller
-. Projections of dorsal tubercles sharply pointed. With a ventri-anal bearing three
pairs of pre-anal setae ...... Epicrius spinituberculatus sp. n.
Epicrius mollis (Kramer), 1876.
Gamasus mollis Kramer, P. Arch. Naturgesch. 1876, 42 : 82 (fig.) ; Berlese, A., Redia, 1916,
12 : 150. Epicrius mollis, Tragardh, I., Ark. Zool. 1942, 34a (4) : 3 (fig.).
Epicrius geometricus Canestrini, G. & Fanzago, F. Atti 1st. Venet. 1877, 1 : 60 ; Haller, G.
Arch. Naturgesch. 1881, 47 : 190 ; Canestrini, G. Prosp. Acarof. Ital. 1877, (5) 4 : 131
(fig.) ; Berlese, A., A. M.S., Padova, 1886, 30 : 8 (fig.) ; George, C. F., Naturalist, 1906 : 265
(fig.)-
The writer follows Berlese (1916) and Tragardh (1942) in considering G. mollis
to be the nymphal (deutonymphal!) stage of E. geometricus. The two main objec-
tions to the acceptance of this synonymy are that Kramer's description and figure
are not sufficiently detailed for the specific identity of the species and that the
original figure of mollis shows only one pair of dorso-lateral protuberances whereas
geometricus is figured with two pairs. It is probable, however, that Berlese examined
the type material of geometricus before coming to any decision regarding the syno-
nymy of the species so it is assumed that the original figure of geometricus is
inaccurate. Berlese (1886) figured geometricus with one pair of dorso-lateral pro-
tuberances and certainly no Epicriid with two pairs of these structures has been
described since Canestrini & Fanzago (1887).
The following re-description of E. mollis is based on material collected over a period
of twelve months from litter under a Spruce stand in South Bedfordshire. 1
Dorsal shield. The dorsal surface of the larva is incompletely covered by a weakly
sclerotized shield bearing nine pairs of setae distributed as in Text-fig. 1. These
setae are probably homologous with those occuring on the anterior shield of the larvae
of the free-living Laelaptoidea (Evans, 1953). The interscutal membrane posterior
to the shield is granulated and bears three pairs of long whip-like setae. In the pro-
tonymph the dorsum is completely covered by a weakly sclerotized shield (Text-fig. 2) .
The chief features of interest in the chaetotaxy of the shield are :
(1) the addition of five pairs of dorsal setae (D) making a total of ten pairs ; and
(2) the addition of setae ni in the anterior half of the shield.
The relative lengths of the setae Di to D5 are approximately the same in both
stages, i.e. D2 and D4 are considerably shorter than Di, and D3. Setae D6 and D7
in the protonymph are much shorter than D7, D9 and Dio. The writer has been
unable to distinguish, with certainty, between larval and protonymphal setae in the
posterior half of the shield. The sclerotized dorsum of the deutonymph is charac-
terized by the development of a pair of large dorso-lateral protuberances (Text-fig. 3).
1 Collected during investigations financed by H. M. Forestry Commission whilst the writer was on the
staff of Rothamsted Experimental Station, Harpenden, Herts.
176
A REVISION OF THE FAMILY EPICRIIDAE
A REVISION OF THE FAMILY EPICRIIDAE
177
The relative lengths of the setae Di to Dio are approximately the same as in the
protonymph. The deutonymphal setae in the anterior half of the shield are indicated
by the symbol n2. There is no increase in the number of setae on the dorsal shield
Text-figs. 4, 6. Epicrius mollis (Kr.). Fig. 4, dorsum of female. Fig. 6, venter of
female. Abbreviations as Fig. 1.
from deutonymph to adult (Text-fig. 4). The dorsum of the adult (both sexes) is
strongly sclerotized and characteristically ornamented. The ornamentation com-
prises bi- and trifurcate tubercles forming a polygonal network (PI. 1). The dorsal
setae are setose (Text-fig. 5). The chaetotactic pattern is shown in the figure.
Setae D8 which are considerably shorter than Do, in the deutonymph are almost
ZOOL. 3, 4.
I4§
178 A REVISION OF THE FAMILY EPICRIIDAE
equal in length to them in the adult. The inornate region surrounding the dorso-
lateral protuberance carries two setae and a large pore-like structure.
Ventral shields. The ventral surface of the larva is weakly sclerotized. The
sternal shield, extending from the posterior margin of coxae I to the posterior margin
of coxae III bears three pairs of simple setae. The oval anal shield is provided
with a pair of para-anals and a post-anal seta. These setae are long and extend
beyond the posterior margin of the body. The interscutal membrane posterior to
coxae IV has four pairs of setae of which two pairs are situated between the sternal and
anal shields. The sternal and anal region in the protonymph is similar to that in the
preceding stage. The region between the sternal and anal shields is provided with
three pairs of setae. The sternal shield in the deutonymph bears four pairs of setae
and is more strongly sclerotized than in the larva and protonymph. The anal
shield has three long setae.
The venter of the female is shown in Text-fig. 6. The first pair of sternal setae are
situated on small platelets (jugularia) surrounded by striated membrane. Sternal
setae II and III stand on a distinct shield situated between coxae II and III and the
metasternals on separate shields postero-lateral to it. The separation of setae IV
from the main sternal shield is difficult to detect in specimens not cleared in lactic
acid. The sternal shield is not fused with the endopodals. The geniti- ventral shield
is large and flask-shaped, and bears two pairs of setae. The epigynal portion of the
shield is poorly developed being reduced to a narrow semi-transparent band along
the anterior margin of the shield. The region between the geniti-ventral and the
ventri-anal shields has a pair of simple setae. The ventri-anal bears a pair of pre-
anals and para-anals, and a post-anal seta. The para-anals and the post-anal seta
are situated behind the anal opening. There is a pair of conspicuous pore-like
structures on the lateral margin of the shield. The remaining sclerotized structures
of the venter are a bi-furcate tritosternum, a porose plate situated posterior to coxae
IV and the ventro-lateral extension of the dorsal shield, which is provided with a
network of tubercles and a number of setae. The interscutal membrane in the region
of the jugularia and lateral to the coxae is richly provided with short spines.
The vental surface of the male differs considerably from that in the female (Text-
fig. 7). The jugularia are present in normal position, but the region between coxae II
and IV is occupied by a well-sclerotized shield bearing four pairs of setae and three pairs
of " pores/' This shield, the sterniti-geniti- ventral, extends beyond the posterior
margin of coxae IV. It is truncated anteriorly but strongly convex posteriorly.
It is not fused with the endopodals. The male genital orifice is situated between
coxae III and is covered by two plates. A pair of setae protrude from under the
postero-lateral margin of the anterior plate. The ventri-anal shield is fused with
dorsal shield, but the setae associated with the anal region are present in the same
position as in the female. The interscutal membrane posterior to the sterniti-
geniti- ventral shield carries three pairs of setae.
Gnathosoma, pedipalps and chelicerae. The ventral surface of the gnathosoma
in the larva is provided with two pairs of setae, namely, the rostrals and the external
posterior rostrals (Text-fig. 8). The corniculi are slender and digitiform. There are
A REVISION OF THE FAMILY EPICRIIDAE
179
four rows of denticles in the region of the ventral groove. The pedipalps comprise
five free segments. The palptibia and tarsus are divided by a distinct suture ven-
trally, but are fused dorsally. The specialized seta on the palptarsus is three-
pronged. The palptrochanter has a spur-like projection ventro-laterally. The
chaetotactic formula for the trochanter, femur and genu is (0-4-5). The ventral
Text-figs. 5, 7, 12, 13. Epicvius mollis (Kr.). Fig. 5, distal end of dorsal seta. Fig. 7,
venter of male. Fig. 12, palptibia and tarsus (ventral) of female. Fig. 13, chelicera of
female.
surface of the gnathosoma in the protonymph bears two additional pairs of setae —
the internal posterior rostrals and the capitular setae (Text-fig. 9). The chaeto-
tactic formula for the palptrochanter, femur and genu is (1-4-5) . In the deutonymph
there is a further increase in the number of setae on the pedipalp. These additional
setae are indicated by the symbol n2 in Text-fig. 10. There is no change in the
chaetotaxy of the pedipalp from deutonymph to adult (Text. fig. n). The structure
180 A REVISION OF THE FAMILY EPICRIIDAE
of the venter of the palptibia and tarsus is shown in Text-fig. 12. These segments
are fused dorsally. The development of the chaetotaxy of the gnathosoma and first
Text-figs. 8, 9, 10, 11. Epicrius mollis (Kr.). Gnathosoma (ventral) of larva (Fig. 8),
protonymph (Fig. 9), deutonymph (Fig. 10) and female (Fig. 11). v., rostral setae ;
ext.p.r., external posterior rostral setae ; int. p. v., internal posterior rostral setae ;
cap., capitular setae.
three free segments of the pedipalp in Epicrius is essentially similar to that described
by Evans (1953) for the genus Typhlodromus Scheuten {Gamasina — Laelaptoidea) .
The tectum in all stages is broadly triangular in shape and has a denticulate
margin (Text-figs. 8-1 1). There appears to be considerable variation in the degree of
denticulation, at least in the series examined.
A REVISION OF THE FAMILY EPICRIIDAE
181
The chelicerae are chelate-dentate.
structure of the chelicera of the female is shown in Text-fig
The dentition is weak in all stages. The
13-
Stigmata and peritr ernes. The stigmata and peritremes are not developed in the
larva. In the protonymph the stigma (st) is situated ventro-laterally in the region
of the third intercoxal space (Text-fig. 14). The peritreme (p) is markedly reduced
and is enclosed with the stigma in an oval depression of the body. The main tracheal
branch is conspicuous. The stigma and peritreme is the same in the deutonymph.
Text-figs. 14, 15. Epicrius mollis (Kr.). Fig. 14, stigmal region of protonymph.
Fig. 15, stigmal region of female, st., stigma ; p., peritreme ; tr., trachea.
In the adult, however, the stigma is enclosed in the heavily sclerotized lateral exten-
sion of the dorsal shield. The stigmal opening is situated on a small protuberence
(Text-fig. 15). The only indication of a peritrematal plate (?) is the presence of a
well-defined line running from the stigma to the level of coxa I and thereby separating
a strip of smooth sclerotized plating from the ornamented dorsal shield.
Legs. All the legs comprise six free segments with the terminal segment incom-
pletely separated into a metatarsus and tarsus. Leg I is the longest in all stages
and the tarsus is without an ambulacral apparatus. In the adult, tibia and tarsus I
are provided with specialized sensory (?) setae ventro-laterally (Text-fig. 16). These
long setae, of which there is one on the tibia and four on the tarsus, are club-like
distally. The swollen head of the seta is minutely setose. The number and position
of these setae are different in the species examined, but since the majority of the
specimens were poorly preserved — the distal end of the setae are easily broken off
in prepared specimens — full use could not be made of this character. The remaining
182
A REVISION OF THE FAMILY EPICRIIDAE
ambulatory appendages terminate in a multi-lobed pulvillus and two claws (Text-
fig. 17). The majority of the setae on the femur to tibia are stout and spinose, and
stand on distinct bases.
~ ^L^L/^ _
2SM
Text-figs. 16, 17. Epicrius mollis (Kr.). Fig. 16, Tibia and tarsus I. Fig. 17,
Ambulacral apparatus of Leg. II.
Text-fig. 37. Berlesiana denticulata sp. n. Tarsus I.
Dimensions.
Larva, 265-280/4 in length and 1 95-200/4 in breadth.
Protonymph, 310-340/4 in length and 245-250/6 in breadth.
Deutonymph, 440-490/4 in length and 305-335/^ in breadth.
Female, 640-660/4 in length and 405-41 o/£ in breadth.
Male, 570-600/4 in length and 360-370/4 in breadth.
A REVISION OF THE FAMILY EPICRIIDAE 183
Locality. This species is the most abundant and widely distributed of the genus.
It has been recorded from moss and forest litter in Italy, Austria, Switzerland,
Germany, Holland, Sweden and the British Isles.
Epicrius canestrinii Haller, 1881
Epicrius canestrinii Haller, G., Arch. Naturgesch. 1881, 47 : 191 (fig.) ; Oudemans, A. C,
Zool. Anz. 1939, 126 : 307.
Epicrius (Diepicrius) parisiensis Berlese, A., Redia, 1916, 1 12 : 152, syn. nov.
Epicrius geometricus, Schweizer, J. Verh. Natf. Ges. Basel, 1922, 33 : 46 $.
The following re-description of E. canestrinii is based on specimens of both sexes
in the British Museum (Nat. Hist). — from the Michael Collection. The specimens
are mounted in Canada Balsam.
Female. The dorsal shield, about 500/^ in length and 350/4 in breadth, shows
the normal network of tubercles. The latter are bi- or trispinate ; the projections
having rounded extremities. The dorsal chaetotaxy is shown in Text fig. 18.
Setae D2 and D4 are considerably shorter than setae D3, but setae D7, D8 and Do,
are approximately equal in length. The dorso-lateral protuberance is well-developed
and broadly triangular in shape. The ornamentation in the region of the protuber-
ance differs from that in the other species of the genus examined by the writer. The
anterior tubercle seta and the dorso-lateral protuberance are distinctly separated
from the posterior tubercle seta by a transverse row of tubercles. The anterior
seta is about twice the length of the posterior seta.
Ventrally, the tritosternum and jugularia are normal for the genus (Text-fig. 19).
Sternal setae II, III and IV are situated on a rectangular shield extending from the
middle of coxae II to the middle of coxae III. The anterior margin of this shield,
in the majority of the specimens examined, is concave and the posterior margin
convex. The geniti- ventral shield is of the usual form with two pairs of simple setae.
The epigynal portion of the shield is well-developed. The interscutal membrane
between the geniti- ventral and anal shields bears four pairs of setae. The four setae
lying between the geniti- ventral and anal shields each lie on a platelet. The number
of platelets appears to be variable. In the female figured by Schweizer (1922) and
which the writer has examined, the two platelets on one side are fused. The anal
shield is small and provided with three setae, of which the para-anals are situated
behind the anal opening.
The gnathosama and pedipalps are normal for the genus. A critical examination
of the chelicerae could not be carried out owing to the state of preservation of the
specimens.
The legs are also normal for the genus. It was not possible to be certain of the
number of clubbed setae on the terminal segments of the first pair of legs. As far
as could be seen three clubbed were present on the tarsus.
Male. The ornamentation and chaetotaxy of the dorsal shield is similar to that
1 Volume 12 of Redia was published on 25.lv. 191 7, but separates of Berlese's paper containing the
description of this species are stated to have been published on 23.viii.1916 (see p. 177).
184
A REVISION OF THE FAMILY EPICRIIDAE
in the female. The chief differences between the sexes is to be seen in the structure
of the ventral surface.
The elongate sterniti-genital shield, posterior to the paired jugularia, has four pairs
of setae and two pairs of " pores." It extends from the middle of coxae II to beyond
the posterior margin of coxae IV. The posterior margin of the shield is truncated
O 100 200/U
19
Text-figs. 18, 19, 20. Epicrius canestrinii Haller. Fig. 18, dorsum of female. Fig.
19, venter of female. Fig. 20, venter of male, ant.tab.s., anterior tubercle seta ;
post, tub.s., posterior tubercle seta.
(Text-fig. 20). The region between the sterniti-genital and anal shields is occupied
by large rectangular shield bearing three pairs of setae. Either side of this shield,
on the interscutal membrane, are two setae. The anal shield, partly fused with
the dorsal shield, is about as broad as long and has three setae and a pair of pore-
like structures. The para-anals lie on the level with the posterior margin of the
anal opening.
Dimensions. Female : 520-550/* in length, 340-352/* in breadth. Male;
480-520/4 in length, 297-320/* in breadth.
A REVISION OF THE FAMILY EPICRIIDAE
185
Locality. The type locality is in the vicinity of Bern, Switzerland. It occurs in
moss and decaying vegetable material and has subsequently been recorded from the
following areas : Meudon (nr. Paris), France (Berlese, 1916) ; Dissenhofen and
Jouxtal, Switzerland (Schweizer, 1922) ; Porth Gwarra, Cornwall, England (Michael
Coll.) ; and Delden, Holland and Sucy-en-Brie, France (Oudemans Coll.).
This species may be readily separated from other species of the genus by the struc-
ture and chaetotaxy of the region of the dorso-lateral protuberance. Haller (1881),
in his original description of the species, deals only with the structure of the dorsum.
He gives a figure of dorso-lateral protuberance and associated setae showing the
separation of the posterior tubercle seta from the tubercle (Text-fig. 21). This
O^ <&<&<& of-
ant.tub.s.
st.tub.s. <&o
/
21
^oj post.tub.s.
Text-figs. 21, 22. Epicrius canestrinii Haller. Ornamentation and chaetotaxy in
the region of the dorso-lateral protuberance of the female. Fig. 21, after Haller
(1881). Fig. 22, specimen in the Michael Collection. Abbreviations as in Fig. 18.
figure agrees very well with the one based on material examined by the writer (Text,
fig. 22).
The male of Epicrius (Diepicrius) parisiensis Berlese agrees in all details with the
male of E. canestrinii.
Epicrius menzeli Schweizer, 1922.
Epicrius menzeli Schweizer, J. Verh. Natf. Ges. Basel, 1922, 33 : 47, figs. $ & $.
The following re-description of E. menzeli is based on the type material kindly
lent to the writer by Dr. J. Schweizer, Birsfelden.
Female. The dorsal shield, 748/6 x 45 1#, is provided with a network of tubercles
as in other species of the genus. The tubercles are bi- and trispinate. The setae
of row D are considerably more uniform in length than in the preceding species —
setae D2 and D4 being approximately equal in length to D3. The dorso-lateral
protuberance is strongly formed and the setae lying immediately anterior and
posterior to it are contained in the same area.
Ventrally, the tritosternum is normal for the genus (Text-fig. 23). The first pair
of sternal setae are situated on distinct platelets. The remainder of the sternal
setae lie on a large sternal shield extending from the middle of coxae II to the middle
i86
A REVISION OF THE FAMILY EPICRIIDAE
of coxae III. Sternal setae II are widely separated from setae III and IV, which lie
in a transverse line along the posterior margin of the shield. A pair of distinct
" pores " lies between setae III and IV. The geniti- ventral shield is large and
flask-shaped, with two pairs of setae. The epigynal portion of the shield extends a
considerable distance beyond the genital orifice. The geniti-ventral is faintly
sculptured and has a pair of " pores." The interscutal membrane between the geniti-
ventral and the anal shields bears seven setae arranged as in the figure. The anal
100
200
24
Text-figs. 23, 24. Epicrius menzeli Schweizer, female. Fig. 24, gnathosoma
(ventral). Fig. 23, venter.
shield has three setae and a pair of large pore-like structures (muscle attachments (?)).
The para-anal setae are situated posterior to the anal opening. The metapodalia
are small, as are the porose-plates posterior to coxae IV.
The gnathosoma and pedipalps are normal for the genus (Text-fig. 24). The
rostrals (240/4), the interior posterior rostrals (7Jfi), the exterior posterior rostrals
(66/i) and the capitular setae (no/^) are distributed as in the figure. The chelicerae
are chelate-dentate and normal for the genus. The legs conform in general with those
in the preceding species. Tibia I (approximately 185/4 in length) is provided with
one clubbed seta and tarsus I (approximately 220/1) with three clubbed setae.
Male, The dorsal shield in this sex measures 627-638/4 in length and 418-429/4
A REVISION OF THE FAMILY EPICRIIDAE
187
in breadth. The ornamentation and chaetotaxy is basically the same as in the
female (Text-fig. 25).
Ventrally, the jugularia are well-developed and widely separated from the sterniti-
genital shield (Text-fig. 26) . The latter has four pairs of simple setae and two pairs
of " pores " distributed as in the figure. The shield is incised anteriorly and truncate
Text-figs. 25, 26. Epicrius menzeli Schweizer, male. Fig. 25, dorsum. Fig. 26, venter.
posteriorly. It is faintly sculptured. The genital orifice is situated between coxae
III. The genital sclerites are as in other species of the genus. The region posterior
to the sterniti-genital is occupied by a large ventri-anal shield bearing thirteen simple
setae and a pair of conspicuous " pores ". The para-anal setae lie posterior to the
anal opening. The shield is faintly sculptured. The porose platelets are situated
postero-lateral to coxae IV. The metapodalia are apparently absent.
188 A REVISION OF THE FAMILY EPICRIIDAE
The gnathosoma and pedipalps are essentially the same as in the female. The
chelicerae were not fully visible in the preparations examined.
Tibia I (176-181/4 in length) is provided with one clubbed seta and tarsus I (178-
198/4 in length) with three clubbed setae.
Dimensions. Female: 748/4 in length, 451/4 in breadth. Male : 627-638/4 in
length, 418-429/^ in breadth.
Locality. The type material comprises one female and three males collected
in damp Beech leaves in a ditch near Bennwil (" Basler Jura "), Switzerland.
Epicrius minor Willmann, 1953
Epicrius (Epicriella) minor Willmann, C, Sitzber. osterr. Akad. Wiss. math-nat. Kl. 1953,
1, 6 : 474, fig. ?.
This species is characterized by the structure of the sternal region of the female,
the only sex known. Sternal setae II lie on the interscutal membrane between the
jugularia and an elongate shield bearing setae III and IV. The geniti- ventral shield
is large and flask-shaped with two pairs of setae and a pair of " pores ". The inter-
scutal membrane between the geniti-ventral and the anal shields has one pair of
setae only. The anal shield bears five setae, of which the paired para-anals are
situated behind the anal opening. The details of the ornamentation and chaetotaxy
of the dorsal shield are not given in the original description of the species.
Dimensions. Female 405/4 in length, 255/4 in breadth.
Locality. " Gipel des Unterberges, etwa 1,300 m., in den Voralpen von Nieder-
osterreich, Waldquadrat in niederem Buchenkrummholz mit zwischenstehenden
jungen Fichten . . . Untervegetation bedeckt den Boden zu einem Drittel, viel
Buchenfallaub. 11.vi.1939, 1 $ " (Willmann, 1953).
Epicrius spinituberculatus sp. n.
Female. The dorsal shield, 594/4 in length and 396/^ in breadth, is richly pro-
vided with tubercles forming a distinct network. The structure of the tubercles is
unique amongst the genus Epicrius in that the distal projections are sharply pointed
and not obtuse as in the other species (Text-fig. 27 and PL II). The number of
dorsal setae is normal for the genus. Setae D2 and D4 are considerably shorter
than D3, whilst D7, D8 and D9 are approximately equal in length (Text-fig. 28).
The dorso-lateral protuberance is well-developed and the setae lying anterior and
posterior to it are contained in the same area.
Ventrally, the tritosternum is normal for the genus (Text-fig. 29). The first pair
of sternal setae are situated on distinct platelets, whilst sternal setae II to IV are on a
sclerotized shield extending from the middle of coxae II to the middle of coxae III.
The distribution of setae II to IV is similar to that in E. menzeli. The geniti-ventral
shield is flask-shaped, being constricted in the region of the anterior of two pairs of
setae which are situated on it. The epigynal portion of the shield is similar to that in
E. mollis. The striated interscutal membrane between the geniti-ventral and anal
shields is provided with two pairs of simple setae. The anal shield is not fused
A REVISION OF THE FAMILY EPICRIIDAE
189
with the dorsal shield and bears three setae. The para-anals are situated behind
the anal opening. A pair of large pore-like structures is present antero-laterally on
the anal shield. The metapodalia and porose plates are present in the positions
shown in the figure.
100
200
1
r©
Text-figs. 27, 28, 29. Epicrius spinituberculatus sp. n., female. Fig. 27, dorsal tubercles.
Fig. 28, dorsal shield. Fig. 29, venter
190
A REVISION OF THE FAMILY EPICRIIDAE
The gnathosoma and pedipalps are normal for the genus. The chelate-dentate
chelicerae were not sufficiently exposed for critical study.
The general structure of the legs is normal for the genus. Tibia I, 187/6 in length,
has one clubbed seta. The number of clubbed setae on tarsus I, which is approxi-
mately equal in length to tibia I, could not be ascertained owing to the state of pre-
servation of the specimen.
rO
50
100
«-200
Text-Fig. 30. Epicrius spinituberculatus sp. n. Venter of male.
Male. The network of tubercles and the relative length of setae D2-D4 on the
dorsum of the male are the same as in the female. The lateral and postero-lateral
setae appear to be relatively longer in length, but this may be due to the distortion
of the dorsal setae in the female as the result of mounting.
Ventrally, the tritosternum and jugularia are normal for the genus. The sterniti-
genital shield bears four pairs of simple setae and two pairs of " pores " (Text-
fig. 30). The genital orifice is situated between coxae III. The region posterior
to coxae IV is occupied by a large ventri-anal shield similar in shape to that in
E. menzeli, but having only three pairs of pre-anal setae. The para-anals are
A REVISION OF THE FAMILY EPICRIIDAE 191
situated posterior to the anal opening. A pair of large pore-like structures is
also present on the shield. The interscutal membrane in the region of the coxae
is richly provided with minute spines.
The gnathosoma and pedipalps are normal for the genus. The chelicerae are
chelate-dentate.
Tibia I, 154/4 in length, is provided with one clubbed seta and tarsus I, 165/4 in
length, has three of the four macro-setae clubbed.
Dimensions : Female : 597/4 in length, 396/4 in breadth. Male : 535-540/4 in
length, 340/4 in breadth.
Locality : A single female (holotype, 1930. 8. 25. 2198) labelled Epicrius canes-
trinii Haller in the Michael Collection in the British Museum (Nat. Hist.) The spec-
imen is without a precise locality, as are the majority of Michaels' specimens.
Also two males (Allotype, 1954.9.8.2 and Paratype 1954.9.8.3), collected by the
writer from a thick layer of humus (F-layer) under bracken in the Leri Valley, near
Dol-y-bont, Cardiganshire, Wales, on 19. iv. 1954.
E. spinituberculatus may be readily separated from other species of the genus by
the structure of the dorsal tubercles, the chaetotactic pattern of the dorsal shield,
and the chaetotaxy of interscutal membrane between the geniti-ventral and anal
shields in the female.
Species Dubiae
The following three species which probably belong to the genus Epicrius, have not
been examined by the writer ; the descriptions are inadequate for their certain
recognition.
Gamasus reticulatus Grube, 1859
Gamasus reticulatus, Grube, A. E., Arch. Naturk. Liv. Ehst.-u. Kurl. 1895, (2) 1 : 459 and
474-
Epicrius reticulatus, Oudemans, A. C, Zool. Anz. 1939, 126 : 306.
The original description is sufficient only to indicate that this species probably
belongs to the genus Epicrius. The ornamentation of the dorsal shield of the unique
specimen is described as " seriebus papillarum fucarum symmetrice reticulato ". The
first pair of legs are without claws. The type locality is in Latvia. Grube's material
may be in Dorpat or Breslau (Oudemans, 1939).
Eugamasus cavemarum Absolon, 1899
Eugamasus cavemarum, Absolon, Ph. C. K., Zool. Anz. 1899, 22 : 324.
Epicrius cavemarum, Oudemans, A. C, Zool. Anz. 1939, 126 : 307.
The description of this species is chiefly based on the male. The ventral surface
in this sex is said to be divided into a number of plates. The sternal plate is small
(or narrow) and rounded off, whilst the genital plate, connected with the anal plate
behind, is prolonged and rounded off anteriorly. Further the " abdominalplatte "
is divided into two distinct plates. The remainder of the description deals with
192 A REVISION OF THE FAMILY EPICRIIDAE
structures which are characteristic of all other species of the genus. The description
of the female is too short to be of value in defining the species.
If Absolon's description of the ventral surface of the male is correct, then E.
cavernarum is undoubtedly a valid species. The writer is inclined, however, to treat
it as a species dubia until the type material can be re-examined.
E. cavernarum was found on the excreta of bats in a cave (Slouperhole) in Moravia,
Czechoslovakia.
Epicrius washingtonianus Berlese, 1916
Epicvius washingtonianus Berlese, A., Redia. 1916, 12 : 151.
Berlese opens his description of this species by stating that " species hanc describere
bene non possum . . . ". The description deals with the ornamentation and chaeto-
taxy of the dorsal shield only and is as follows :
". . . Tamen a caeteris hucusque notis est diversum, quod corniculis dorso-
lateralibus (ad quartos pedes) caret et areolis minus numerosis quam in E. cirrato
gaudet. Pili dorsi (qui adhuc persistunt) breves, nulla barbula ornati, simplices,
forsitan omnes intersese statura subaequales (ad 8o/£ long.). Tuberculi dorsi
plerumque bilobi, rarius trilobi. Ad 6oo/£ long. ; 420/^. lat. Caeteris hucusque
notis specibus maior.
Habitat. Inveni in muscis ad ' Washington ' collectis."
II Genus Berlesiana Turk, 1943
Berlesiana Turk, F. A., Ann. Mag. nat. Hist. 1943, (11), 10 : 855.
This genus was proposed by Turk for Epicrius cirratus Berlese. He considered the
structure of the sternal region in that species to be so distinctive that " when the
types are re-examined a new family will have to be made ". This would appear to
be likely in the light of Tragardh's concept of the family Epicriidae (Tragardh, 1939).
The discovery of a new species undoubtedly congeneric with E. cirratus but with
sternal setae II and III in the female on an undivided shield shows that the structure
of the sternum in the Epicriidae is more variable than Tragardh realised, and
cannot be used as a major character in the classification of the Epicriina.
The ornamentation of the dorsal shield and the structure of the dorsal setae in
E. cirratus separates it from the mollis — group of species. The writer proposes to
retain Berlesiana Turk as a valid genus but at the same time emending the original
definition as follows :
With the general characters of the genus Epicrius but differing from it in
the following characters : tubercles of the dorsum more numerous ; postero-
lateral protuberance poorly developed ; dorsal setae short, stout and strongly
barbed ; geniti- ventral shield in the female more or less parallel-sided ; tarsus
I with less than three pairs of clubbed setae.
Type : Epicrius cirratus Berl., 1916.
A REVISION OF THE FAMILY EPICRIIDAE 193
The genus Berlesiana contains two species which may be separated by the structure
of the sternal shield in female, as follows :
1. Sternal setae II to IV each situated on a separate shield Berlesiana cirrata (Berl.)
-. Sternal setae II and III on an undivided shield . . Berlesiana denticalata sp. n.
Berlesiana cirrata (Berl.), 1916
Epicrius cirratus Berlese, A., Redia. 1916, 12 : 151.
Berlesiana cirratus Turk, F. A., Ann. Mag. nat. Hist. 1943, (n) 10 : 855.
In the original description of this species Berlese mentions the ornamentation
and chaetotaxy of the dorsal shield. The dorsal setae are about 50/* in length.
The dorso-lateral protuberance is stated to be absent, but it is possible that Berlese
may have overlooked it, since a poorly developed protuberance is present in the
normal position in the species described below.
The sternal plate in the female is described as "in partes duas laterales fracto ;
quaeque pars autem fissura transversa plus minusve bene in scutulis duobus rotundis,
in medio piliferis divisa ". The ventri-anal shield is provided with five setae in the
female. The male is not described.
Dimensions. Female : 440/* in length, 300/4 in breadth. Male : " considerably
smaller ".
Locality. This species is known from the type localities only ; the female from
Chianti and Pontedera, and the male from near Genoa. In all localities the species
was found in moss.
Berlesiana denticulata sp. n.
Female. The dorsal shield, 415-425/* in length and 260-265/t in breadth,
completely covers the dorsum of the mite. The ornamentation of the shield con-
sists of numerous tubercles, predominantly tri-lobed (Text-fig. 31). The dorso-
lateral protuberance is not nearly as well developed as in the genus Epicrius. The
dorsal setae with the exception of four pairs of simple setae antero-laterally, are
short (44-55/*), stout and strongly barbed (Text-fig. 32). Setae Di to Dio are
approximately equal in length. The distribution of " pores " is shown in the figure.
Ventrally, the tritosternum and jugularia are as in the genus Epicrius. Sternal
setae II and III are situated on an undivided shield extending from the middle of
coxae II to the middle of coxae III (Text-fig. 33). Setae IV (the metasternals) lie off
this shield. In the holotype the metasternal seta on one side lies on the inter-scutal
membrane but on the other side there are two setae situated on a small shield. This is
undoubtedly an aberration since in the paratypes sternal setae IV are situated on the
interscutal membrane postero-lateral to the shield bearing setae II and III. The
geniti-ventral shield is ornamented with punctations and is roughly rectangular in
shape. Its posterior margin is truncated. The epigynal portion of the shield is
well-developed and denticulate along its anterior margin. The geniti-ventral bears
two pairs of simple setae. The endo-podal plates are poorly developed and not fused
with the sternal shield. The ventri-anal shield is elongate, truncate anteriorly and
194
A REVISION OF THE FAMILY EPICRIIDAE
provided with a pair of pre-anal setae. The surface of the shield is scabrid. There
is a tendency for increased setation of the ventri-anal. In one of the paratypes
there are four pre-anal setae. The ventri-anal is not fused with the dorsal shield
50
100/*
^&^S @D
« d^Sfc^ @D3
s Oi,<?1)Q % ©D5
<b o J
^> ©D9
fclj
Text-figs. 31, 33. Berlesiana denticulata n. sp., female. Fig. 31, dorsal shield.
Fig. 33, venter.
(Text-fig. 34) . A pair of simple setae is present between the geniti- ventral and anal
shields.
The gnathosoma is provided with four pairs of setae ventrally of which the
rostrals are considerably longer than the other three pairs. The pedipalps have
A REVISION OF THE FAMILY EPICRIIDAE
195
five free segments. The chaetotaxy of the palptrochanter, femur and genu is
(2-5-6) . The palptarsus is incompletely separated from the tibia ; the two
segments being fused dorsally. The specialized seta on the palptarsus is three-
Text-figs. 32, 34, 35, 36, 38. Berlesiana denticulata n. sp. Fig. 32, dorsal seta of
female. Fig. 34, postero-ventral region of female. Fig. 35, tectum of female. Fig. 36,
chelicera of female. Fig. 38, venter of male.
pronged. The tectum is denticulate and basically the same as in species of
Epicrius (Text-fig. 35). The chelicerae are chelate-dentate (Text-fig. 36).
Leg I is considerably longer than the body and is without an ambulacral apparatus.
Tibia I, about no/i in length, has no clubbed setae. The chaetotaxy of tarsus I is
shown in Text-fig. 37 (p. 182). This segment has two clubbed setae and, proximally
i 9 6 A REVISION OF THE FAMILY EPICRIIDAE
a long stout seta lying parallel to the longitudinal axis of the segment. Legs II to IV
terminate in a pulvilms and two claws, and are of the form in the genus Epicrius.
Male. The ornamentation and chaetotaxy of the dorsal shield in the male is
essentially similar to that of the female.
Ventrally, the jugularia are relatively large and are distinct from the sterniti-
genital shield which extends from the middle of coxae II to well beyond the posterior
margin of coxae IV. This compound shield bears four pairs of simple setae distri-
buted as in Text-fig. 38. The genital orifice is situated between coxae III and its
structure is normal for the family. The surface of the shield appears to be corrugated
in preserved specimens. It is not strongly sclerotized. The ventri-anal shield is
similar in form to that in the female except that it bears an additional pair of pre-anal
setae. The striated interscutal membrane between the sterniti-genital has a pair
of simple setae. There is also evidence of increased setation of the ventral shields
in the male. In one specimen two short setae were present posterior to the last
pair of " sternal " setae on the sterniti-genital.
The gnathosoma, pedipalps and legs are basically the same as in the female.
Dimensions. Female : 415-425^ in length, 260-265^ in breadth. Male : 400-
405/^ in length, 240-250/^ in breadth.
Locality. Five females and seven males from litter under Beech at Boxhill,
Surrey (collected by Messrs. S. K. Eltringham and K. H. Hyatt, 21.viii.1953).
Holotype female, 1954 . 9 . 8 . 9 ; Allotype male, 1954. 9. 8. 10; Paratypes, 1954.9.8.
11-20.
GEOGRAPHICAL DISTRIBUTION OF THE EPICRIIDAE
Any discussion on the zoogeography of the free-living Acarina is open to criticism
on the basis that very little work has been done on them outside the Palaearctic and
Nearctic Regions. It is, therefore, not possible to be dogmatic about the limits of
distribution of any group of free-living mites. Bearing this in mind, it can be stated
that, at present, the Epicriidae are recorded only from the Palaearctic and Nearctic
Regions ; ten " species " from the former and only one species from the latter. 1 Epi-
crius mollis (Kr.) appears to be the most widely distributed species in the Palaearctic
Region having been recorded from a number of localities between Sicily in the south
and Sweden in the north.
The writer has been unable to find representatives of this family in recent collec-
tions of free-living mites from Uganda, India, Singapore and Rennell Is.
DISCUSSION
The examination of the above mentioned species of the genera Epicrius and Berle-
siana enables one to discuss in some detail the value of the morphological characters
which have formed the bases of the various classifications of the Epicriidae put
toward by previous workers. In several of these investigations, particularly those
1 Baker & Wharton (1952) state that " on Guam Epicrius sp. was found among the rhizomes of epi-
phitic ferns ..." in which case this is the first record of the family outside the Palaearctic and Nearctic
Regions.
A REVISION OF THE FAMILY EPICRIIDAE 197
of Tragardh, on the comparative morphology of the Mesostigmata, the classification
has been built up on the evidence supplied by the critical examination of only one
species, Epicrius mollis (Kr.). The following outline of the development of the
classification of the family since its erection by Berlese (1885) will serve as the basis
for discussion.
Berlese (1885) considered the genus Epicrius and Podocinium Berl. to constitute a
a subfamily (!) Epicriidae of the family Gamasidae. The characteristic features of
this subfamily were the elongate first pair of ambulatory appendages and the orna-
mentation of the body of the mites. Epicrius and Podocinium were separated on the
degree of development of the peritreme. Later (Berlese, 1892), Epicrius was placed
with Zercon Koch and Seiodes Berl. in the Zerconidae, apparently on account of the
position of the genital orifice in the male. This classification was retained by Berlese
(1913) in his introduction to the " Acarotheca Italica," except that the Zerconidae
was divided into the Zerconini and the Epicriini. Vitzthum (1929, 1931 and 1941)
followed, in the main, Berlese's classification and at the same time emphasized the
close relationship between the Epicriidae and the Zerconidae.
Tragardh (1938), in the first of his important contributions to the classification
of the Mesostigmata, separated the Epicriidae and the Zerconidae ; placing the former
in the Sejina (= Liroaspina) and the latter in the Gamasina on differences in the
structure of the genital and sternal regions in the females. The Epicriidae was
regarded as a primitive group because of the absence of a distinct epigynal shield and
the segmentation of the sternal region. The Sejidae ( = Liroaspidae) and Epicriidae
were separated as follows :
"A. Sternal shields III and IV fused, forming a narrow transverse shield,
separated from the remaining sternal shield .... Fam. Sejidae.
A A. All sternal shields fused ...... Fam. Epicriidae."
In 1946 (19460:) the same general classification of the Liroaspina was given except
that the sternal region of the Epicriidae was now described as " sternal shields I free,
shields II-IV fused." But after examining the structure of the genital region in
the male, the Epicriidae was removed from the Liroaspina to the Epicriina (Tra-
gardh, 1946&). At the same time the Epicriina was stated to resemble the Zer-
conina in this character. Therefore, the final result of Tragardh's researches were
to separate the Epicriidae and the Zerconidae into distinct cohorts, chiefly on the struc-
ture of the genital region in the females.
The writer disagreed with Tragardh's interpretation of the structure of the genital
region in the female of the Epicriidae and could find no evidence for his statement
that an epigynal shield is not developed in this group (Evans, 1955). The geniti-
ventral shield in both Epicrius and Berlesiana are basically the same as that in the
Laelaptoidea. This is shown in Text-figs. 39-41, where a comparison is made between
the genital region of two species of Laelaptoid mites and E. mollis. The writer
therefore, suggested the inclusion of the Epicriidae and Zerconidae in the Epicriina,
following Vitzthum (1941).
The second important morphological character used by Tragardh in the classifi-
cation of the Liroaspina and Epicriina is the structure of the sternal shield in the
198 A REVISION OF THE FAMILY EPICRIIDAE
females. It is on the basis of the degree of fragmentation of the sternal shield in
the female that the Epicriidae was originally separated from the Liroaspidae and,
later, the Liroaspidae from the Epicrosejidae (Tragardh, 1952). This separation of
the families appears to have been made after the examination of one species from each
family. From the present revision of the Epicriidae it is quite obvious that the frag-
mentation of the sternal region in this group shows considerable variety and can no
•P- 39
g-v.sh.
Text-figs. 39-41. Lateral view of the sterniti-geniti- ventral region of the females of
two species of Laelaptoid mites (Figs. 39 and 40) and Epicrias mollis (Kr.), (Fig.
41). ep., epigynal portion of the geniti-ventral shield ; g-v-sh., geniti-ventral shield.
longer be used as a major character in the classification of the family. The following
types of sternal " shields " were encountered in the females :
1. Sternal setae II free ; III and IV on a shield, (E. minor).
2. Sternal setae II and III on a shield ; IV free, (E. mollis, B. denticulata).
3. Sternal setae II, III and IV on a shield, (E. canestrinii, E. menzeli..
E. spinituberculatus) .
4. Sternal setae II, III and IV each on separate shield, (B. cirrata).
Jugularia are present in both sexes in all species.
A REVISION OF THE FAMILY EPICRIIDAE 199
The generic concept in the Epicriidae also has the structure of the sternal region
in the females as its basis. In the recent classification given by Baker & Wharton
(1952) the genera Epicrius and Berlesiana are placed in separate families ; the former
in the Epicriidae and the latter in the Liroaspidae. In the present classification
both genera have been included in the Epicriidae and the subgenera Diepicrius
and Epicriella have been relegated to the synonymy.
SUMMARY
1. A revision is given of the family Epicriidae with a discussion on the classifica-
tion of the Epicrhna.
2. Two genera, Epicrius and Berlesiana, are considered valid. The former
contains five valid species and three of uncertain status, and the latter two species.
3. Keys to species are given for the identification of both sexes of Epicrius and the
females only of Berlesiana. Epicrius {Diepicrius) parisiensis Berl. is considered to be
a synonym of Epicrius canestrini Haller.
4. The following two species are described as new : Epicrius spinituberculatus
and Berlesiana denticulata.
ACKNOWLEDGMENTS
The writer is extremely grateful to Dr. Jos. Schweizer for the loan of material
from his collection, to Dr. L. van der Hammen for specimens from the Oudemans
Collection, to Dr. G. Lombardini for comparing specimens with those in the Berlese
Collection, and especially to Dr. H. W. Parker for his criticisms of the manuscript.
The photographs are the work of Mr. M. G. Sawyers.
REFERENCES
Absolon, Ph. C. K. (1899). Uber die fauna der Hohlen des mahrischen Devonkalkes (Vor-
laufige Mittheilung) . Zool. Ariz. 22, 324.
Baker, E. W. & Wharton, G. W. (1952). An Introduction to Acarology. Macmillan.
465 pp.
Berlese, A. (1882-1892). Acari, Myriopoda et Scorpiones hucusque in Italia reperta. Padova.
(1885). Acarorum Systematis. Bull. Soc. ent. Ital. 17, 129.
(1913)- Acarotheca Italica, Systema — Acarorum Genera. Firenze.
(1916a). Centuria Prima di Acari Nuovi. Redia, 12, 32.
(19166). Centuria Seconda di Acari Nuovi. Redia, 12, 150.
Canestrini, G. & Fanzago, F. (1886). Intorno agli Acari Italiani. Atti. 1st. Venet. (5) 4,
131.
— ■ — (1885). Prospetto dell' Acarofauna Italiana. Padova, 60.
Evans, G. O. (1953) • Some mites of the genus Typhlodromus Scheuten, 1857 from S.E.
Asia. Ann. Mag. nat. Hist. (12), 7, 615.
( I 955). A collection of Mesostigmatid mites from Alaska. Bull. Brit. Mus. {nat. Hist.)
Zool. 2, 9 ; 287-307.
George, C. F. (1916). Lincolnshire Mites, Epicrius. Naturalist, 265.
Grube, A. E. (1859). Verzeichniss der Arachnoiden Liv-, Kur- und Ehstlands. Arch.
Naturk. Liv.-Ehst.-u. Kurl. 1, 459 and 474.
Haller, G. (1881). Acarinologisches. Arch. Naturgesch. 47, 190.
Kramer, P. (1876). Zur Naturgeschichte einiger Gattungen aus Familie der Gamasiden.
Arch. Naturgesch. 42, 82.
200 A REVISION OF THE FAMILY EPICRIIDAE
Oudemans, A. C. (1939). Neue Funde auf dem Gebiete der Systematik und der Nomenklatur
der Acari V. Zool. Anz. 126, 306.
Schweizer. Jos. (1922). Beitrag zur Kenntnis der terrestrischen Milbenfauna der Schweiz.
Vevhandl. der Naturf. Ges. Basel, 33, 47.
Tragardh, I. (1910). Acariden aus dem Sarekgebirge. Naturw. Under such, des Sarekge-
birges, 4, 432.
(1938). Further contributions towards the comparative morphology and classification
of the Mesostigmata. Ent. Tidskr. 59, 123-158.
(1942). Zur Kenntnis der Gattung Epicrius Berlese. Ark. Zool. 34a, 4, 3.
(1946a). Outlines of a new classification of the Mesostigmata based on comparative
morphological data. Lunds Univ. Arsskr. 42, 1-37.
(19466). Contributions towards the comparative morphology of the Mesostigmata
(Acarina) VII. Ent. Tidskr. 67, 88-108.
(1952). Acarina collected by the Mangarevan expedition to South Eastern Polynesia . . .
(Mesostigmata). Ark. Zool. (2) 4, 46-90.
Turk, F. A. (1943). Studies of Acari. — I. The British Species of Epicriidac, with Descriptions
of a new Genus and two new Species. Ann. Mag. nat. Hist. (11), 10, 855-860.
Vitzthum, H. G. (1929). Die Tierwelt Mitteleuropas, VII Acari. Leipsig.
(1931)- Kiikenthals Handbuch der Zoologie, 3, 2, 143.
(1941)- Acarina in Bronn's Tierreich 5, Abt. 4, Buch 5, 778.
Willmann, C. (1953). Neue Milben aus den ostlichen Alpen. Sitzber. dsterr. Akad Wiss.
math.-nat., Kl., 1, 6, 474.
EXPLANATION OF PLATES.
PLATE 1.
Ornamentation of the dorsal shield in Epicrius mollis (Kramer), x 1000.
PLATE 2.
Ornamentation of the dorsal shield in Epicrius spinituberculatus sp. n. x 800.
'
PR ■ TED
2 SEP 1955
<^
PRESENTED
2 SEP 1955
PRINTED IN GREAT BRITAIN BY
ADLARD AND SON, LIMITED,
BARTHOLOMEW PRESS, DORKING.
PRESENTED
1 uah 1956 THE MONK SEALS
(Genus Monachus)
JUDITH E. KING
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 3 No. 5
LONDON : 1956
THE MONK SEALS
(GENUS MONACHUS)
BY
JUDITH E. KING
Pp. 201-256 ; Pis. 3-8 ; 12 Text-figures
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 3 No. 5
LONDON: 1956
THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, is
issued in five series, corresponding to the Departments
of the Museum, and an Historical series.
Parts will appear at irregular intervals as they
become ready. Volumes will contain about three or
four hundred pages, and will not necessarily be
completed within one calendar year.
This paper is Vol. 3 , No. 5 of the Zoological series.
PRINTED BY ORDER OF THE TRUSTEES OF
THE BRITISH MUSEUM
Issued January, 1956 Price Eighteen Shillings
THE MONK SEALS (Genus Monachus)
By JUDITH E. KING
CONTENTS
Page
Synopsis ............ 204
Synonymy ............ 204
M . monachus ........... 205
History ........... 205
Distribution ........... 206
Taxonomy ........... 209
Description . . . . . . . . . . . 210
M. tropicalis ........... 215
History . . . . . . . . . . .215
Distribution . . . . . . . . . . .216
Taxonomy ........... 218
Description . . . . . . . . . . . 218
M. schauinslandi ........... 220
History ........... 220
Distribution ........... 222
Description ........... 223
Specimens Examined .......... 223
Osteology ....... .... 226
Comparison of skulls . . . . . . . . .226
Dentition ........... 232
Supernumerary bones . . .. . . . . .236
Scapula . . . . . . . . . . . 237
Humerus ........... 239
Radius and ulna .......... 239
Manus ............ 239
Pelvis and sacrum . . . . . . . . 239
Femur . . . . . . . . . . .241
Patella . . . . . . . . . . .241
Tibia and fibula . . . . . . . . . .241
Pes . . . . . . . . . . . .241
Vertebral column .......... 242
Ribs ............ 245
Sternum . . . . . . . . . . 245
Growth ............ 246
Relationships . . . . . . . . . . .251
Acknowledgments .......... 252
Bibliography ........... 252
zool. 3, 5.
204
THE MONK SEALS
SYNOPSIS
An account is given of the history, distribution, features and habits of the three species of
monk seal Monachus monachus, M. tropicalis and M. schauinslandi.
A list is given of the skulls, skeletons, skins and stuffed animals that have been examined.
The skull, dentition and skeleton are described and compared with those of other Phocidae.
The members of the genus Monachus are shown to be much more nearly allied to the Southern
than the Northern Phocids.
The relative growth rates of the various parts of the skull are given, and show that the facial
region grows more quickly than the cranial region.
The genus Monachus — the monk seals — consists of only three species. M. monachus
(Hermann 1779) lives in the Mediterranean area, M. tropicalis (Gray 1850) in the
West Indies, and M. schauinslandi Matschie 1905 in the Hawaiian Islands. They
are interesting animals, but because of their widely scattered distribution,
remoteness and scarcity, not a great deal is known about them. In this paper an
attempt will be made to bring together as much information about the monk seals
as it has been possible to find.
SYNONYMY
Genus MONACHUS Fleming, 1822
Monachus Fleming, 1822, Philos. Zool. 2 : 187 (footnote). Phoca monachus Hermann.
Pelagios F. Cuvier, 1824, Mem. Mus. H.N. Paris, 11 : 196. Phoca monachus Hermann.
Pelagius F. Cuvier, 1826, Diet. Set. Nat. Paris, 39 : 550. For Pelagios.
Pelagus McMurtrie, 1834, Cuvier's Animal Kingdom, 71. For Pelagios.
Pelagias Gray, 1837, Mag. Nat. Hist., N.s. 1 : 582. For Pelagios.
Pelagocyon Gloger, 1841, Gemeinn. Naturgesch. 1, 34 : 163. Phoca monachus Hermann.
Rigoon Gistel, 1848, Nat. Thier, fur hohere Schulen. X. New name for Pelagios F. Cuvier.
Heliophoca Gray, 1854, Ann. Mag. Nat. Hist. 13 : 201. Heliophoca atlantica Gray = Phoca
monachus Hermann.
I. Monachus monachus (Hermann, 1779)
Phoca monachus Hermann, 1779, Besch&f. Berlin Ges. Naturf. Freunde, 4 : 501, pis. 12, 13.
Phoca albiventer Boddaert, 1785, Elench. Anim. 1 : 170.
Phoca bicolor Shaw, 1800, General Zoology. 1,2: 254.
Phoca leucogaster Peron & Lesuer, 18 16, Voy. aux Terres Austr. 2 : 47 (footnote).
Phoca hermanni Lesson, 1828, Diet. Class. Hist. Nat. 13 : 416.
Monachus mediterraneus Nilsson, 1838, K. Svenska Vet. Ah. Handl. 1837 : 238.
Phoca crinita Menis, 1848, II Mare Adriatico, 153.
Heliophoca atlantica Gray, 1854, Ann. Mag. Nat. Hist. 13 : 202.
II. Monachus tropicalis (Gray, 1850)
Phoca tropicalis Gray, 1850, Cat. Mamm. in colln. Brit. Mus. part 2. Seals, 28.
wilkianus Gosse, 1851, A Naturalist's Sojourn in Jamaica, 308 (footnote).
III. Monachus schauinslandi Matschie, 1905
Monachus schauinslandi Matschie, 1905, Sitzungs-Berichte der Gesellschaft Nat. Freunde, Berlin,
258.
THE MONK SEALS 205
I. Monachus monachus
History
The monk seal appears to have been always well known to the inhabitants of the
Mediterranean area. Clark (1952) notes that bones of this seal were found in upper
palaeolithic levels at Grimaldi, and there is evidence that seal hunting was quite an
important activity in classical Greece. The skins were collected partly for the
superstitions surrounding them and partly for use as clothes by the poor fisher folk.
Boots, tents and dresses made of skin were said to protect one from lightning ; a
seal skin drawn round a field and then hung up by the door would save the field from
hailstones, and the right flipper was supposed to be a cure for insomnia when put
under the head at night. Greek writers such as Plutarch, Pliny, Homer and
Aristotle knew and wrote of the seal. Aristotle must have examined one with care
as he gives quite an accurate account in the Historia Animalium, but Keller (1887)
says that on the whole the poets found the animal horrible and deformed, and
objected to its oily smell. The presence of seals gave rise to many stories. The
" half animal fisheaters " on the west coast of Africa were said to have made a pact
with the seals not to interfere with each other's fishing, and in the Odyssey a woman
who died on board ship was thrown overboard to serve as food for the seals and
fishes. Because of their love of sun and sea the seals were put under the protection
of Phoebus Apollo, the sun god, and Poseidon, the sea god. The seal was frequently
shown alive in these times. Even then their docility and intelligence were noted,
and Pliny tells how he saw some which answered by growls when their names were
called. In earlier days there must have been more monk seals round the Greek
coasts than there are now, as several towns have taken the name of the seal. Phocis
is the name of an ancient Greek district that stretched past Mount Parnassus to the
Gulf of Corinth, and there is at the present time Foca at the north end of the Izmir
Gulf, Turkey, and Foca, an inland town in Yugoslavia. Keller (1887) notes that
very ancient pre-Darian coins (ca. 500 B.C.) show a picture of a seal, and that coins
from Rhodes show a seal's head. He also mentions that etymologically the Greek
work for " phoca " means the swollen or plump animal, and must have been first
applied, though not in a taxonomic sense, to the monk seal.
Some of the later history has been summarized by Monod (1932) . An Italian map
of the fourteenth century shows an island between Lanzarote and Fuerteventura in
the Canary Islands called Ya de Uegi marini — the island of sea wolves — to-day
called more simply Lobos Island ; and in 1341 Niccoloso di Recco included seals in
an inventory of the Canary Islands. In the fifteenth century Portuguese explorers
found numerous bands of monk seals and killed many for their skins, and in 1434
Alfonso Baldaya started the industrial exploitation of the seals in the bay Rio del
Oro, north of Cap Blanc. A traveller named Zarco reached Madeira in 1418 and
named a small cove Camara de Lobos because of the seals he found there.
Rondoletius published his Aquatilium Historia in 1554 and included a drawing and a
short Latin description of " de Vitulo maris mediterranei ", but the first specific name
was given by Hermann (1779) who described an animal from the Dalmatian coast
2 o6 THE MONK SEALS
and called it Phoca monachus. Buffon (1782) mentions that seals from the Adriatic
were kept in captivity in France and Germany in 1760, and describes a female seal
that was on exhibition in Nimes in 1777. He also gives a detailed description of a
male seal that was caught on 28th October, 1777, " off the island of Guarnero " in
the Adriatic. He saw this seal in Paris in December, 1778, and from the similarity
of his description of it and of its locality and habits it seems that it must have been
the same animal described by Hermann — a fact which Cuvier noted as early as 1813.
Hermann saw the seal in Strasbourg at the end of October and beginning of
November, probably in 1778, and says that it was being taken on tour by a company
of Venetians who were going to Paris to show it to the King. He mentions that the
seal was said to have been captured in the autumn of 1777 in the Dalmatian Sea at
Ossero. Ossero is on the island of Cherso in the Gulf of Quarnero, off the Dalmatian
coast. Buffon notes that the seal died in August, 1779.
Since that time there have been many references, though mostly of the occurrence
of single individuals. Cuvier (1813) describes a seal that was kept in captivity for
two years in a very small bath, only a foot longer and two feet wider than the animal
itself. In this remarkably small enclosure it spent 9-10 hours of each day in six
inches of water that were drained off each night. The London Zoological Gardens
has three times kept a monk seal for short periods. The first specimen, a young
female, arrived in May, 1882, but died the same day ; the second was in 1894, when
a young animal from Madeira lived there for three and a half months ; and the last
was in 1910, when a second animal from Madeira lived for four months. In 1926
the American Museum of Natural History received skins and skeletons of three seals
from the Desertas Islands ; these were believed to be the first Mediterranean seals
in American museums. More recent references have again been concerned mainly
with isolated occurrences, and there is at the time of writing a male monk seal from
near Oran that has been in captivity for about two years in the Jardin des Plantes
in Paris.
Distribution and abundance
The Mediterranean monk seal is known from the shores and islands of the
Mediterranean and the western coast of North Africa (figs. 1 and 2). It has been
recorded from : Gulf of Almeria, Spain ; Cabrera, Balearic Is. ; Toulon ; Corsica ;
Cape Teulada and the Gulf of Cagliari in Sardinia ; the island of Pelagosa in the
Adriatic ; the Gulf of Quarnero and Fort Opus in Yugoslavia along the Dalmatian
coast ; the Gulf of Salonika in north Greece ; Cape Caliacra, and generally in the
Black Sea and Bosphorus ; Tantoura ; El Arish ; Port Said ; the island of Galite
off the Tunisian coast ; Oran ; Madeira and the Desert a Grande Islands ; the
Canary Islands, and along the African coast, including Cap Barbas, Baie d'Etoile
and Baie du Levrier, to Cap Blanc.
Budker (1945) says that the southern limit along the African coast is 20 49' N.
(approximately the latitude of Cap Blanc), that its limit of distribution is influenced
by the temperature of the sea and corresponds with the 20 C. winter isotherm. In
the British Museum collections there is an Ascarid from a monk seal taken in Senegal
1
208
THE MONK SEALS
(approximate latitude 15 N.). It has been in the Museum since 1863, but there is
no other information about the host.
Apart from the few references given by Monod (1932) for the fourteenth and
fifteenth centuries there do not seem to be any other accounts of large scale
commercial exploitation of the monk seal, although during a visit to Madeira in
Fig. 2. — Map of Mauritanean coast and Atlantic islands, and inset of Cap Blanc to show
distribution of M. monachus.
1945, Cadogan (1945) heard that " in 1943-4 a consignment of some twenty-five
seal pelts had been seized by the Customs Officials at Funchal, and it was not thought
likely that any further attempt would be made to commercialize them for the
present ". Any commercial use of an animal necessarily implies its abundance in
the area. Admiral W. H. Smyth, writing of the period 1810-1824 (in Flower, 1932)
says that " between Alexandria and Benghazi , . , we found fish and seals in
THE MONK SEALS 209
abundance ", but from then up to the present day there have been references only
to single animals or small groups. Barcelo (1875) said that seals were very common
on the shores of the Balearic Islands at the time he wrote, but Cabrera writing in
1914 notes their disappearance from these parts. Aharoni (1930) notes that the
seal is quite often offered for sale by fishermen from Askalon and Jaffa. Bertram
(1943) says that a seal was seen off El Arish in about 1941, and according to Monod
(1945), Agacino (1950) and Postel (1950) there must still be reasonably large colonies
along the western coast of Africa down to Cap Blanc. Sixty seals were seen by
Postel just north of Cap Blanc, and twenty-one by Agacino along the coast of Rio de
Oro, and Cousteau and Dumas (1953) note the presence of a herd of about two
hundred seals at Port Etienne. An adult female was surprised and killed in a cave
in Corsica in 1947 (Troitzky, 1953), and Butler (in litt.) saw a seal in the Gulf of
Salonika in 1950. There is still a small breeding colony in the more isolated regions
of the Desertas Islands, although the fishermen there regard it as an enemy to their
livelihood. A recent report in The Times (nth June, 1954) says that the monk seal
still appears yearly on the Turkish coast. The main stronghold of the monk seal at
present seems to be along the coast of Rio de Oro, and if not molested it is probable
that its numbers will be maintained there.
Taxonomy
The first specific description of the monk seal is that given by Hermann in 1779.
He named as Phoca monachus a male animal captured at Ossero, on the island of
Cherso, off the Dalmatian coast in 1777, and seen by him in Strasbourg. He gives
a very detailed description of the seal and its habits, and also a drawing of the
whole animal with details of the head and flippers (PL 4, a). In 1782 Buffon, not
knowing of Hermann's work, described the same seal which was, in 1778, on show
in Paris. He gives a very good drawing of the animal (PL 4, b) but only refers to it
as " Le phoque a ventre blanc ". These two descriptions form the basis of
practically all the succeeding names which have been applied to the monk seal.
Boddaert, in 1785, used Buffon's paper as the basis for his very brief Latin
description of Phoca albiv enter, a specific name which was in use for many years.
In volume two of the third edition of Pennant's Quadrupeds, published in 1793,
Pennant gives a description of the Pied Seal, mentioning Buffon, but adding " This
I saw at Chester ; it was taken near that city in May, 1766 ". It is difficult to
know whether Pennant was applying Buffon's description to a seal actually taken
near Chester, Cheshire, in which case it was very unlikely to have been a monk seal,
or whether he was confusing the name with Cherso on the island of that name in the
Adriatic. The drawing he gives is not very good, (PL 4, c) and shows an animal
with a broad white ring round its neck and a white spot behind one flipper, the rest
of the body being black. Pennant also describes the Mediterranean seal, quoting
Hermann, but giving no picture. Shaw in 1800 uses Pennant's Pied Seal and
Buffon's Phoque a ventre blanc for his Phoca bicolor, and used Phoca monachus for
the Mediterranean seal. He uses the drawings previously given by Buffon and
Pennant and labels both of them " Pied Seal var,"
210 THE MONK SEALS
In a brief footnote, in which he quotes a passage from Buff on, Peron (1816) uses
the name Phoca leucogaster, and in another equally brief footnote Fleming (1822)
first suggests the use of the generic name Monachus, " Some seals, as Ph. monachus,
are said to have four incisors in each jaw. Such will probably be constituted into a
new genus, under the title Monachus ". In 1824 Cuvier suggested Pelagios as a
new generic name for Phoca monachus Hermann, a name which has given rise to
many variants ; Cuvier himself in 1826 called it Pelagius, McMurtrie (1834) used
Pelagus, and Gray (1837) use d Pelagias. Lesson in 1828 renamed Hermann's
species Phoca hermanni, and Nilsson in 1838 called it Monachus mediterraneus. In
1 841 Gloger called the monk seal of the Mediterranean Pelagocyon monachus, and in
1848 Menis used the name Phoca crinita for an animal presumably from the Adriatic
(book not seen). Gistel, also in 1848, proposed the generic name Rigoon instead of
Pelagios of Cuvier. Giebel (1848) linked the monk seals with the southern Phocids
under the genus Leptonyx and the subgenus Leptorhynchus . This part of his
classification is given here in full :
" Leptonyx
a. Stenorhynchus
1. L. serridens
2. L. leopardinus
b. Leptorhynchus
3. L. weddellii
4. L. rossii
5. L. monachus."
The last synonym was given by Gray in 1854, when he named the new genus and
species Heliophoca atlantica on the skin and skull of a young animal from Deserta
Grande, Madeira. The skull and stuffed skin of this animal are now in the British
Museum collections (Reg. No. 1853. 10.6.4, 1063a.)
Description
Fully grown adults are about 8-9 ft. long. Gavard (1927) gives the length of a
female which had produced a pup as 2-42 m. (7-9 ft.), but does not say whether
this length included the hind flippers or not ; Troitzky (1953) gives the nose to tail
length of an old female as 278 m. (9*1 ft.), and both Postel (1950) and Agacino
(1950) note that the biggest animals that they saw were about 3 m. (9*8 ft.) long.
Monod (1945) gives the length of a male animal as 2*9 m. (9-5 ft.). Of the two
adult skins in the Museum collection the length of the male is 2*38 m. (7.8 ft.) and
of the female 2 • 1 m. (6 -9 ft.). These measurements are from nose to tip of tail, but
should be regarded with caution as that of the male was taken from a rather crumpled
dressed skin, and that of the female from an undressed but folded skin. The weight
of the female seal measured by Gavard was 300 kilos (661 lb.), and that measured by
Troitzky was 302 kilos (666 lb.) without the viscera.
There is a certain amount of variation in the colour of the adult seal. Gavard
says that it is all black except for some patches, particularly a large one of a dirty
THE MONK SEALS 211
white colour round the navel, and some little yellowish patches on top of the head.
Agacino (1950) notes that the seal is spotted ventrally, but that there is some
variation in colour and the old male may be a silvery colour all over. The light
yellowish beige colour of the female described by Troitzky was also probably due to
age. Cousteau and Dumas (1953) saw a large white bull in one of the caves on the
islands of La Galite, but this may have been the silvery colour of old age. The monk
seal now in the Jardin des Plantes in Paris is thought to be an adult animal and is
chocolate brown dorsally shading to greyish fawn ventrally. It did not appear to
have a white ventral patch but had several whitish scar marks along the back. An
examination was made of the few skins of this species in the British Museum
collections. The dressed skin of a fairly young male animal, length from nose to tip
of tail 1 -75 m. (5*74 ft.) (1894.7.27.3, 1063b), is dark blackish brown dorsally,
with a slightly yellowish appearance due to the yellow tips of the dark brown hairs.
This shades to light brownish yellow ventrally, but without a light ventral patch.
The skin of an adult male (length 2-38 m., 1890. 12. 30.1) is generally dark blackish
brown, slightly yellowish along the centre of the back and belly due to the yellow
hair tips. In the centre of the belly, slightly nearer the fore flippers than the tail,
is a roughly diamond-shaped patch of dirty yellow colour, about 73 cm. long and 58
cm. wide. The whole of the skin, particularly under the chin, the sides of the neck
and the centre and hinder region of the back, is covered with irregularly placed
streaks and spots of yellowish hair. These marks show on the under surface of the
prepared skin and may possibly be due to scars. The skin of the adult female
(length 2-i m, 1894.7.27.2, io63g) is more like the young animal than the adult
male. It is dark blackish yellow dorsally, the hairs being dark brown with yellowish
tips. This shades to light greyish yellow ventrally, and there is no light ventral
patch. The back shows a few light scar-like streaks similar to those found on the
male. These whitish streaks were also noticed on the seals examined by Hermann
and Carruccio (1893) ; they are found on both sexes. The white ventral patch
seems to be irregular in its occurrence.
The hairs of the adult are very short and bristly and lie close to the body. They
are approximately half a centimetre long. The appearance of the young seal
before it moults its natal coat is described from the youngest skin in the Museum
collections (1892. 11. 7.1, 1063I). The length of the skin from nose to tip of tail is
1 -4 m. (4.4 ft.). Dorsally it is a rich dark brown, shading at the level of the fore
flippers to a lightish brown ventrally. On the belly, slightly nearer the fore flippers
than the tail, is a roughly diamond-shaped patch of a dirty yellowish colour with a
few very small light brown spots on it. The patch is approximately 34 cm. long
and 28 cm. wide at its widest point. There is a small light brown area along the
upper lip, and the whiskers are also light brown and oval in cross section. The
texture of the hair of this young seal is quite different from that of the adult. It is
soft and woolly, the hairs are 1-1J cm. long and do not lie close to the body as they
do in the adult.
The whiskers range in colour from light yellow to brown ; they are smooth, not
wavy as in P. vitulina, and oval in cross section. Nails are present on both fore and
212 THE MONK SEALS
hind flippers. That on the first digit of the fore flipper is about 2-54 cm. (1 in.)
long, and the others decrease slightly in size towards the fifth digit. The nails on
the hind flippers are very small and inconspicuous. The tongue has a notch in its
anterior end.
Lobstein (18 17) described in some detail the anatomy of the viscera of a female
seal that died in Strasbourg in 1815 after touring France and Germany for two years
and Troitzky (1953) gives a brief description of the viscera of a full term foetus, but
the most detailed recent investigation into the internal anatomy of M. monachus
was done by Dieuzeide (1927) on a young male seal that was captured near Oran in
December 1926, and which lived for a few months in the Experimental Station at
Castiglione, Italy. A brief summary of the results, taken from Dieuzeide unless
otherwise noted, is given here for completeness. The length of the male animal was
1 -68 m. (5-5 ft.) from nose to tip of tail, and of Lobstein's female 2-13 m. (7-0 ft.).
The oesophagus which lay mainly to the left of the trachea and large blood vessels
was 60 cm. long and 4 cm. in diameter and was very dilatable. The oesophagus of
the adult female (Lobstein) was 97 cm. long, the small intestine 14-5 m. and the
large intestine 1-3 m. The whole alimentary canal was 17-5 m. (5-74 ft.) — eight
times the length of the animal. There was no definite line of demarcation between
the oesophagus and the stomach, which looked more like a dilatation of the
oesophagus. The height of the stomach was about 25 cm., its width about 12 cm.
and it led into the small intestine, which had a length of 12 m. and a diameter of
3-3 J cm. The caecum was a small pocket and the large intestine measured a metre
in length by 4 cm. in diameter. The liver was large, measuring 40 x 40 cm. and
consisted of six long pointed lobes. There was a large venous sinus (a dilatation of
the inferior vena cava) and the round ligament (the remains of the umbilical vein)
was well represented. The gall bladder was multilobulate and when full measured
13 x 13 mm. The left lung was the larger, weighing 950 gm., while the right weighed
850 gm. The kidneys were two oval masses, the right anterior to the left, with a
combined weight of 1 kg. 140 gm. They were lobed and had an extensive vascular
network. The left renal vein was large and of practically the same diameter as the
vena cava. It was formed from the union of three large vessels and the superficial
plexus of the kidney. On the right side the vena cava followed the inner border of
the kidney and received vessels from it. The right kidney was supplied from the
aorta by two renal arteries which arose a little anterior to the one going to the left
kidney. The suprarenals were small and almond shaped, measuring 33 X 14 mm.
The bladder was very thick (9 mm.) and measured 15 x 4 cm. The prostate was
reduced and the testes were internal, measuring 4x1.5 cm. The penis was 8 cm.
long and the baculum 7.8 cm. long. The heart was oval in shape, and Dieuzeide
gives a detailed description of its anatomy.
Several species of parasites have been recorded from the stomach and intestine
(Joyeux and Baer 1936, Baylis 1937, Markowski 1952). There are two Nematodes —
Contracaecum osculatum, (Rudolphi 1802) and Porrocaecum decipiens (Krabbe 1878),
and four species of Cestode — Diphyllobothrium coniceps Linstow 1907, D. elegans
(Krabbe 1865), D. lanceolatum (Krabbe 1865) and D - hians (Diesing 1850).
THE MONK SEALS 213
Remarkably little information is available about the breeding habits of this seal.
Practically all that is known is in a recent paper by Troitzky (1953). In this she
mentions that after a gestation period of eleven months the pups are born on land in
September and October and are fed by the mother for six or seven weeks. The
female has four teats on the posterior part of the abdomen and lies on her side when
feeding the pup. At the end of this time the pup moults its woolly coat and enters
the water for the first time. Troitzky says that the young seals stay with their
mothers for three years, that they do not begin to breed until they are four years
old, and that mating of the adult animals takes place about seven or eight weeks
after the birth of the pup. The sum of eleven months' gestation plus seven or eight
weeks before mating makes a total breeding cycle of thirteen months, and although
it is usual in most Phocids for the female to bear a pup at the same time every year,
Troitzky says that this seal only has a pup every alternate year.
Apart from Troitzky's paper, most of the information about breeding times has to
be inferred from records of still born pups, foetuses and young animals. Dathe
(1934) records that a young female seal was caught on 19th September, 1933, on the
Dalmatian coast. The umbilicus was not yet healed and the animal was thought
to be only a few days old. Its length was approximately 90 cm. and it weighed 26
kilos. This little animal was kept in captivity and was going to be taken to
Frankfurt Zoological Gardens. It was fed by bottle six or seven times a day on a
mixture of half gruel and half milk with a little cod liver oil and freshly rubbed fish
paste. On 26th September it was taken to Split, preparatory to moving to
Frankfurt, but the journey re-opened a wound on its stomach, caused probably by
a fish hook, and on 29th September it died, its length then being 1 . 20 m.
Carruccio (1893) notes that a foetus 50 cm. long was taken from a pregnant female
on 21st May, 1891, and Gavard (1927) mentions a captive female that produced a
still-born pup on 14th April, 1926. This pup weighed 2.25 kilos and measured 62
cm. Both these foetuses are of such a size that they would have been full term and
born about September, and it is strange that Postel (1950) should say that the pups
are born in the spring. Agacino (1950), who went to Las Cuevecillas, Rio do Oro on
26th December, 1945, says that at that time the smallest seals were 1.5 m. long,
and that a mother was seen to be feeding her pup. This must have been a pup
born very late in the season, probably about the middle of November.
The voice is said to be a sharp strong cry from the bottom of the throat (Cuvier,
1813), while Agacino (1950) says that when they are annoyed they make a noise like
a wounded dog. Hermann said that the one he observed had a voice like that of a
hoarse dog and that sometimes it would howl. This seal could not tolerate dogs,
and would try to drive them away by clapping its teeth. The seals kept in captivity
have all been noticeably intelligent and docile animals. They have become attached
to their keeper and would recognize him, follow him about, and even obey his orders
to a certain extent.
The feeding habits, as observed in captivity, are very interesting. The animal
described by Hermann ate about 14 pounds of fish daily and in order to stress the
expense of keeping it, inquisitive spectators were told that it ate only the best fish,
214 THE MONK SEALS
such as eels and trout. It did occasionally receive eels and carp, if paid for by the
spectators, but usually it had whiting. It took the fish either out of the keeper's
hands or caught them in its tank, but preferred to eat them in water. It seized the
fish by the head, squeezed and shook them a few times and then swallowed them
whole. Often intestines of the fish were found in the water, and although the
keeper thought this was done deliberately, Hermann suggested that the insides of
the fish came out accidentally when it was squeezed. Buff on saw the same animal
while in Paris and said that there it was fed mainly on carp and eels, preferring the
latter. The fish were sprinked with salt, the eels eaten whole, but the carp were
crushed with the teeth, let fall and then the belly of the fish ripped open and the
entrails removed. The fish was then seized by the head and swallowed. Cuvier
also notes that the entrails were removed and the fish swallowed head first. The
seal at present in Paris was also seen to swallow its fish head first, and this has also
been observed in Halichoerus grypus and Phoca vitulina. Indeed it seems possible
that this method might be the normal one for Pinnipeds generally, as it would avoid
any injury to the seal by the backwardly projecting fins, scales and spines of the fish.
Two female seals described by Gavard (1927) also disembowelled their fish unless
they were very small, and also were unable to pick the fish off the ground, and could
eat them only in water. They ate sardines, bonito and octopus — about 12 kg. a
day. Boettger (1951) notes that native fishermen along the African coast say that
the seals eat fish and lobsters (Palinurus) , and remains of fish of the genera Dentex
and Labrax have been found in the stomach of a seal captured off Sardinia (Carruccio
1893). A monk seal in the Gulf of Salonika was seen to be playing with a large fish,
tossing it into the air and catching it again (Butler in litt.)
Troitzky (1953) describes an adult female seal that was killed in a cave on Corsica
in September, 1947. It was found to be pregnant and a full-term foetus was
removed, but could not be revived. The pup, a female, was 120 cm. long and
weighed 17 kilos. It was dark, greyish black colour, with a white ventral patch.
Troitzky notes that the pup in its colouring did not differ from descriptions of what
she regarded as " typical " M. monachus, but says that the mother was not so.
She observes that in its dentition, its great size, and the time of breeding the adult
resembles M. monachus, but the shape of its head with a long snout, the light colour,
the absence of the white ventral patch, and the second digit of the fore flipper longer
than the first are characters not associated with that species. After reviewing other
members of the Phocidae she comes to the conclusion that these characters are more
like those of Arctic Phocids, and says that the most logical conclusion is that this
female is a hybrid, the result of a cross between a monk seal and, probably, Phoca
groenlandica, and that it is interesting that such a hybrid should have been able to
produce a pup.
It is considered extremely unlikely that such a cross could have taken place.
Phoca groenlandica and other members of the Arctic Phocidae have not been recorded
from the Mediterranean in Recent times, and indeed, they seldom occur south of
Arctic latitudes. From the description of the teeth — worn, broken and diseased —
it is assumed that the animal was old, and the light colouring was probably also due
THE MONK SEALS
215
to age. The white ventral patch is not of universal occurrence amongst the monk
seals. It is not possible to comment on the other two characters, from lack of exact
information, but from the photograph of the adult seal, as far as it is possible to see
the shape of the fore flipper does not appear to be unusual and the drawings and
photograph of the skull do not appear to differ in any way from undoubted skulls of
M . monachus in the Museum collection.
II. Monachus tropicalis
History
The first reference to the monk seal of the West Indies is that given in the account
of the second voyage of Columbus. At the end of August, 1494, the ship anchored
by the rocky island of Alta Vela, south of Haiti (= Hispaniola), and the men that
went ashore killed eight " sea wolves " that lay sleeping on the sand (Kerr, 1824).
The next record chronologically, and the first for Florida, is that of Herrera who,
while describing Ponce de Leon's discovery of the Dry Tortugas Islands (lat. 24 10'
N. long. 83 55' W.) on 21st June, 1513, said that a foraging shore party took
fourteen seals (Moore 1953). Du Tertre (1667) was told by Brother Charles Poncet,
who had been to Guadeloupe, that he had seen at least twenty asleep under the trees
near the shore, and many of them were killed. Dampier (1705) noted that there
were seals on the Alacrane Islands in 1675, Sloane (1707) saw them on the Bahama
Islands in 1687, and Olafsen (1774) makes a reference to the seal of the Antilles.
Hill (1843) gives a description of a young seal from Pedro Kays, and Gosse (1851)
published an account of a voyage in 1846 by Mr. George Wilkie to Pedro Kays,
where he saw several seals and killed a few. It was on this voyage that the type
skin of Phoca tropicalis was obtained. In 1883 the U.S. National Museum received
a mounted skull and skin from a female seal captured off Cuba earlier in the same
year and presented by Prof. Felipe Poey (True and Lucas, 1885). In 1886 H. L.
Ward and Prof. F. Ferrari Perez of the Mexican Geographical and Exploring Survey
set out from Campeche to the Triangles to search for Monachus. (Ward, 1887b).
Although they were only on the islands from 1st to 4th December, forty two
specimens were taken away and shared between the two members of the expedition.
Of the specimens retained by H. L. Ward a complete articulated specimen (1887 . 8 .
5.1), and a skin and skull (1889. n. 5.1) are now in the British Museum (Natural
History), and an articulated specimen (899c) is in the Cambridge Zoological Museum.
These last two specimens were purchased from Ward by F. D. Godman of Cambridge.
A recently born pup was taken back to Campeche, but it lived there only a week.
A female seal was captured at the Triangles in 1897 and lived for nearly 5 J years in
New York Aquarium, where it died in 1903. (Anon 1903). E. W. Nelson and E. A.
Goldman, during their biological investigations of Mexico from 1892 to 1906
(Goldman, 1951), spent the period 18th to 23rd June, 1900, on the Triangles. Their
main object there was to obtain specimens of seals, but apart from remarking that
" in quest of these animals we were very successful ", no mention is made of their
abundance, or how many were killed. The New York Aquarium received four more
216 THE MONK SEALS
seals — an adult male and three yearlings — in June, 1909 (Townsend 1909). These
were obtained from a dealer in Yucatan who presumably got them from from the
Triangles or the Alacrane Islands. Several seals were seen on the Tortugas Islands
during the period 1903-8 and two were kept in a moat for some time, where they
became fairly tame (Moore, 1953). Six seals were captured by a fishing vessel in
1915 and taken to Pensacola, where they were kept in captivity for some time and
then turned loose, when bathers in the area objected to their presence (Allen 1942).
Townsend (1923) notes that a seal was killed near Key West, Florida in March, 1922.
Gunter (1947) gives sight records of seals along the Texas coast in 1926 and 1932,
and Lewis (1948) says that a young seal was killed at South West Kay in the Pedro
Group in 1939. A. C. Wheeler (in litt.) saw two seals on the beach of Drunken
Man's Cay, about two miles south of Kingston, Jamaica in November, 1949.
Distribution and abundance
The West Indian monk seal was at one time abundant in the Gulf of Mexico and
off the islands in the Caribbean Sea (Fig. 3.). The presence of numerous Seal Cays
and Islands and Lobos Cays show how widely the seal was distributed in the area.
1. Seal Cay, south of Long Island, Bahamas, lat. 22 38' N., long. 75 ° 54' W.
2. Seal Cay, south of Caicos Bank, Bahamas lat. 21 10' N, long. 71 47' W.
3. Seal Island, north of Anguilla, Leeward Is, W. Indies lat. 18 24' N., long.
63°2o'W.
4. Cay Lobos, north of Cuba, lat. 22 25' N., long. yy° 36' W.
and Allen (1887a) gives the following :
5. Seal Keys — on the coast of Honduras in about lat. 16 N., a few miles north-
east of the Mosquito Coast.
6. Seal Key — about 200 miles further south along the same coast in about
lat. 12 40' N.
It has been recorded from the Bahama Islands ; Key West, Florida ; Galveston
Bay and Brazos, Texas ; the Triangle Islands to the west of Yucatan, lat. 20 55' N.
long. 92 12' W. : the Alacrane reef to the north of Yucatan, lat. 22 32' N., long.
89 45' W. ; the shores of Cuba and Jamaica ; the Pedro Kays to the south of
Jamaica, lat. 17 N., long. 77 30' W. ; Alta Vela, a rocky island south of Haiti ;
and Guadeloupe.
That the seal has formerly been abundant is evident from some of the earlier
accounts. Dampier (1705) said of his visit to the Alacranes in 1675 : " Here are
many seals . . . the Spaniards do often come hither to make Oyl of their Fat ;
upon which account it has been visited by English-men from Jamaica, particularly
by Capt. Long : who having the Command of a small Bark, came hither purposely
to make Seal-Oyl." Sloane (1707) gives the following account of the seals of the
Bahamas, " The Bahama Islands are fuTd with Seals, sometimes Fishers will catch
one hundred in a night. They try or melt them, and bring off their Oil for Lamps
to these Islands." In 1856 a Mr. Alexander was on the Triangles and saw only two
THE MONK SEALS
217
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218 THE MONK SEALS
living seals, but remains of skeletons and hides indicated a once flourishing business ;
and although H. L. Ward does not say exactly how many he saw in 1886, there must
have been quite a large colony as he killed over forty animals (Ward, 18876). Allen
(1887a) suggested that at the time of his writing seals were still present on the islets
of Salt Key Bank, north of Cuba, the isles off Yucatan, and probably the isles
between. In 1897 at the time of the capture of the seal for New York Aquarium
about thirty were observed on the Triangles (Anon 1903), but as late as January, 1911,
according to Lewis (1948) about two hundred seals were killed in this locality and at
the time his paper was published he thought that there were perhaps still a few left
there and on the South West Kay in the Pedro Group. In view of the fact that
seals have been seen as recently as 1949 near Jamaica, and that Moore (1953) quotes
a " well informed and responsible friend " who knows of the whereabouts of seals
somewhere within their former range, it seems likely that a remnant of this species
is still living.
Taxonomy
During Mr. George Wilkie's visit to the Pedro Kays in 1846 (Gosse, 1851) he
obtained the skin of a monk seal which he gave to Gosse. Gosse sent this skin, which
had no bones with it, to the British Museum where J. E. Gray published a description
of it under the name Phoca tropicalis Gray, 1850. This skin has been stuffed and is
in the Museum collection (Reg. No. 1847.2.2.2). Gosse (1851) republished the
description of the seal given by Hill (1843), and in a footnote on p. 308 suggested
the specific epithet " wilkianus ", but he gave no generic name, and moreover, was
already antedated by Gray. In 1866 in the Catalogue of the Seals and Whales in the
British Museum Gray repeated the descriptive paragraph he gave in 1850, but used
the name Monachus tropicalis.
Description
The nose to tail length of an adult male animal is between 7 and 8 ft., females being
in general slightly smaller. Townsend (1906) gives the length of an old female with
very worn teeth as 9 ft., but he was probably measuring to the tips of the hind
flippers. The nose to tail lengths of both the skeleton in the British Museum,
(probably a female) and that in Cambridge, (sex unknown) are both about 7 ft. 3 in.
The nose to tail length of a dressed skin of an adult male (1889. 11. 5.1, 1064b) is
7 ft. 5 in., and Ward (18876) gives nose to tail lengths of two pregnant females as 7 ft.
1 in. and 6 ft. 6 in. A female measuring 6 ft. 11 in. to the tip of the hind flippers
and which had been in New York Aquarium for 5^ years weighed 360 pounds at
death. This seal died from fatty degeneration of the heart, liver and kidneys, so
presumably a healthy animal of this size would weigh less.
Ward (18876) gives the following description of the colour of the adult seal :
" Adults are grayish brown or grisled on the back, a result of the Vandyke-brown
hairs being tipped with light horn-color, the lower surface ochreous yellow to
yellowish white. Females seem to have much less of the yellow or white on the
ventral surface. From the black pelage of the extremely young to that of the adult
THE MONK SEALS
219
there is an intermediate stage of yellowish gray on the dorsal surface, shading to
almost a perfect ochre on the ventral portions." This is the most reliable account of
the colour, and corresponds well with that of the stuffed specimen in the Museum,
which is the type of Phoca tropicalis Gray. This animal is dark brown, and slightly
lighter on the sides and belly. The hairs of the back are dark brown with a lighter
tip, while those of the sides and belly have a more yellow tip. The hairs lie close to
the body and are extremely short, the longest — those on the sides — being about 1 cm.
long. A dressed skin of an adult male from the Triangles in the Museum collection
(1889. 11. 5. 1, 1064b) is dark blackish brown all over, with a slight yellowish tinge
due to the yellow tips of the hairs. New born pups are black in colour and the hair
is long, soft and woolly.
The whiskers are yellowish horn colour, some being slightly darker at their bases.
Those of the Museum specimens are oval in cross section. Nails of appreciable size
are present only on the fore flippers. That on the first digit is about an inch long,
those on the remaining four digits decreasing gradually in size. The nails of the
hind flipper are very small.
The description of the eye is given by Ward (18876) : " The pupil is medium
sized, round, and well defined, the iris is light reddish brown in color, and with but
little of the sclerotic coat showing. Over the cornea there appears a deadening
film . . .", which he attributes to the strong reflection of the tropical sun from the
coral sands.
Young animals are born about the beginning of December. Ward (18876) was on
the Triangles from 1st to 4th December and killed five females with full-term
foetuses, and noticed another female with a new-born pup. The female has four
teats. One of the foetuses measured 85 cm. from nose to tip of tail, was covered in
black woolly hair and had black whiskers. Measurements of the skull and skeleton
of this pup as well as those of adult animals collected by Ward are given by Allen
(1887a).
Ward examined the stomach contents of several animals, but found only fluids
and large numbers of intestinal parasites several inches in length. Gosse (1851)
recorded the opinion of the " more experienced fishermen ", who said that the seals
fed " as generally on molluscous animals as on fish ", but there is no proof of this
The animal noted by Hill (1843) lived for four months in captivity, without eating
and when it died " the fat was four inches thick and yielded four gallons of oil "
The skull of this animal, which was then, as Hill (1846) says " an undescribed Seal "
was exhibited at a meeting of the Zoological Society of London in September, 1846
Unfortunately it has not been possible to trace this skull, which was probably the
first specimen of the West Indian monk seal to reach this country, arriving here
shortly before the skin sent by Gosse which became the type.
The West Indian monk seal appears to be a fairly noisy animal. Hill (1843) said
that his young animal " grunted, barked, growled and snarled like a dog ", and
Ward (18876) said that the voice of the young was " a long drawn out guttural ' ah '
with a series of vocal hitches during its enunciation ". Townsend (1909) also noted
that it was noisy, and the young often roared harshly.
220 THE MONK SEALS
On land at least the seal seems to be rather lethargic. Ward (18876) notes that
unless the seals were approached closer than three or four feet they showed no
interest or alarm. Closer than that they would rouse themselves, bark, and move
off a little. When Ward and his party attacked a group of seals they got more
excited and would make savage rushes, and would then fall back on their dead
fellows and bite and shake them. Nevertheless, as Ward notes " the whole aspect
of the animals was one of indecision . . . they only roused themselves to action
on being individually attacked ". This behaviour is not peculiar to the monk seal,
but seems to be common to all seals. Ward also notes the peculiar circumstance
that several of the animals he collected had a growth of minute algae upon their
backs and flippers, especially the hinder ones, so that they appeared quite green.
Although Ward says they are neither curious nor playful, Hill (1843) notes that
the young specimen he kept in captivity was lively, and those kept in New York
Aquarium were certainly playful. The two which were received in 1897 had the
habit of filling their cheeks with water and squirting it at visitors, while the seal
which arrived in 1909 amused itself by tossing flipperfuls of water into the faces
of visitors.
III. Monachus schauinslandi
History
The Hawaiian or Sandwich Islands are a chain of small islands near the centre of
the North Pacific Ocean between 18 55' and 28 25' N., and 154 48' and 178 25' W.
(Text-fig. 4) ; Honolulu on the island of Oahu being 2,100 miles S.W. from San
Francisco and 3,445 miles S.E. from Yokohama. The chain stretches 1,578 miles
from E.S.E. to W.N.W. All the islands are uninhabited, except Midway which is a
transpacific cable station and sea plane base administered by the U.S. Navy, and
Niihau, Kauai, Oahu, Molokai, Maui and Hawaii. The entire chain forms the
Hawaiian Islands Bird Reservation.
Perhaps owing to the remoteness of the islands on which they live, there are very
few references to the Laysan monk seal. In the early part of the nineteenth century
seals must have been numerous as Bryan (1915) records that in 1824 the brig
" Ainoa " set out from Hawaii on a sealing voyage in that area, and in 1859 * ne
" Gambia " returned to Honolulu with 1,500 seal skins and 240 barrels of seal oil,
some of which was probably from Midway Id., which was discovered on this trip.
In 1893 a Mr. J. J. Williams visited Laysan and heard of an earlier expedition that
had killed sixty or seventy seals on the island (Atkinson and Bryan, 1913). In 1905
Matschie published a description of a seal skull brought back from Laysan by Dr.
H. Schauinsland, and named it after him. The U.S. revenue cutter " Thetis "
returned in 191 2 after a cruise to Midway and Laysan and brought back a seal skin
which was presented to the Bishop Museum in Honolulu (Bryan, 1915) and parts of
three others which are in The U.S. National Museum. Thirty five seals were seen
on Pearl and Hermes Reef in 1913 (Atkinson and Bryan 1913), and Dr. Wetmore,
who visited the area with the U.S.S. " Tanager " Expedition in 1923 saw a number of
seals and collected ten for the U.S. National Museum (Bailey, 1952). In 1940 about
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222 THE MONK SEALS
half a dozen were seen round Midway (Blackman, 1941) and records of visits to the
islands in 1949 and 1951 show that quite large numbers were seen resting on the
beaches. The first, and it is believed, the only seal of this species to be kept in captivity
outside the Hawaiian Islands was a young animal which was presented to San Diego
Zoo by the Honolulu Zoo in 1951 (Anon, 1951). It was captured on French Frigates
Shoal but only lived about three months in captivity.
Distribution and abundance
The Laysan seal has been recorded from the following of the Hawaiian islands :
Ocean Island, Midway Island, Pearl and Hermes Reef, Lisiansky Island, Laysan
Island, French Frigates Shoal, and a stray animal has been recorded from the coast
of Hawaii, although they do not generally appear to go so far eastwards.
In 1824, x ^59 an d 1893 the seals were obviously very numerous, although in
Schauinsland's own account of his visit to Laysan (Schauinsland, 1899) ne OIU y
mentions that " seals come singly, indeed very seldom by the island ". In 1912,
Elschner (1915) who was also on the "Thetis" during its trip, noted that there were
many seals on French Frigates Shoal and Pearl and Hermes Reef. Also on the latter
island thirty five seals were seen in 1913 (Atkinson and Bryan, 1913). The U.S.S.
" Tanager " Expedition sailed from Honolulu in April, 1923, for a visit to the Hawaiian
Islands. The Bulletin of the Bishop Museum in Honolulu (Bull. 10, 1924) only
mentions that several seals were seen on Lisiansky on this voyage, and that two
skins and skulls were collected ; Allen (1942) however, notes that Dr. A. Wetmore
while on this expedition saw colonies of seals on Pearl and Hermes Reef and on Ocean
Island, and estimated the total population to be about four hundred, and Bailey
(1952) notes that he brought back ten specimens for The U.S. National Museum.
Sixty eight seals were seen on Pearl and Hermes Reef in 1930, and five on Laysan in
1936 (Bailey, 1952), and Blackman (1941) who stayed for six months on Midway in
1939-40 saw about six seals. More recent visits to the islands (Bailey, 1952) show
that the animals are still fairly numerous there. About thirty were seen on Laysan
and over a hundred on Midway. A count of seals made in 1951 showed a total of
407 on the beaches, the largest populations being on Laysan (119) and Pearl and
Hermes Reef (180). The others were seen on French Frigates Shoal, Maro Reef,
Lisiansky and Midway. It was suggested that the large numbers on the beaches
could perhaps be accounted for by the presence of tiger sharks in the water.
The monk seals are distinctive in being the only truly warm water form of Phocid,
with the exception of the northern elephant seal (Mirounga angustirostris) . So far
as the distribution of the three species is concerned it is not difficult to accept the
occupation of the West Indian islands from a source in the Mediterranean and along
the Mauritanian coast, as the Canary Current passing down the latter coast would
bear the emigrants to the eastern limit of the North Equatorial Current sweeping due
west to the Caribbean Sea. The extension of the range of the genus to Hawaii and
across the Isthmus of Panama is feasible when it is accepted that Phocids are capable
of considerable overland journeys. For example Hayes (1928, p. 106) records that
Captain Scott, when in the Antarctic, found seal carcases as far as fifty miles inland
THE MONK SEALS 223
and 5,000 ft. above the sea. The Isthmus of Panama at its narrowest is much less
than fifty miles, and its lowest height above sea level less than 200 ft. If rivers
were exploited by the seals the distances travelled overland might have been still
further diminished. The North Equatorial Current in the Pacific could well have
borne the animals to the islands they now occupy. Allen (1942) suggests that the
colonization of the Pacific by the monk seal was from the West Indian stock in
Tertiary time, when there was a waterway connecting the Atlantic and Pacific.
Description
Of the few records available only those of Matschie and Bailey (1952) give any
information about the external features and Bailey includes some excellent photo-
graphs. A male, probably not full grown, collected by the " Thetis " measured 5-7
ft. from tip of nose to tip of tail, and a female 7-5 ft. Matschie gives the following
description : "A scalp which I have for examination has very short, thick, bristly
hair, loam coloured, with a silky shine. The lips are yellow-gray, and it is somewhat
brownish on the front side of the neck. Most of the whiskers are light horn coloured,
a few dark brown, and all with very thin points. The sides of the body of the stuffed
animal in Bremen Museum are lighter than the back, the breast and stomach are
whitish." Bailey notes that the underparts are light straw yellow and the back is
dark slate grey.
While on the " Tanager " Expedition Dr. Wetmore examined the stomachs of
seals for parasites. All the seals had abundant nematodes in the stomach and
Chapin (1925) has described these as a new species Contracaecum turgidum.
Cestodes, Diphyllobothrium Mans, were also found.
The birth of the young seals evidently takes in January. The " Thetis " saw
pups at this time in 1912, and one was born during the visit of Governor Frear to
Pearl and Hermes Reef in January, 1913 (Atkinson and Bryan, 1913). The young
male seal that lived in San Diego Zoo for a short time (Anon, 1951) was received in
May and died probably at the beginning of September. It was therefore about eight
months old at the time of its death. It is described as a nursing pup of approximately
three feet long. It had a silver-tinged coat which was dark brown above, paler on the
sides and nearly white ventrally. The muzzle was whitish and had many coarse
whiskers.
Atkinson and Bryan note that the seals are fearless and readily handled, and
Blackman (1941) says that if cornered they threaten the intruder by opening their
mouths widely and uttering an abrupt barking noise. He also says that they do
not migrate, and probably feed on squid and fish. Bailey notes that an animal
collected by Henry Palmer in 1891 had its stomach full of half digested fish ; and he
also makes an interesting observation that a large male seen in 1949 had a greenish-
coloured face, but what the cause of this was he did not say.
SPECIMENS EXAMINED
The following specimens of Monachus have been examined. The list includes
stuffed animals and skins, and unless otherwise mentioned the specimens are in the
British Museum (Natural History) .
224 THE MONK SEALS
M. monachus
Skulls only
i. 1853.10.6.4,1063a. Presented by R. Macandrew.
Deserta Grande Is. Madeira.
Type of Heliophoca atlantica Gray, 1854.
Skull and lower jaw of immature animal. Skull with the dorsal part of the
cranium, inter orbital and nasal regions missing.
Stuffed animal also in the collection.
2. 1063b. Presented by R. Macandrew.
Deserta Grande Is. Madeira.
Skull incomplete — maxillary region bearing teeth, and fragmentary lower jaw
only.
3. 1934.8.5.4 Collected by Barrett-Hamilton.
Mediterranean.
Complete skull and lower jaw.
4. 1951 . 4 . 17 . 1. From Rothschild Collection, Tring.
Skull and lower jaw. Zygomatic arches broken.
Skulls and skeletons
1. 1063c. Presented by M. Verreaux.
Algiers.
Complete skull and skeleton of an immature animal.
2. 1863.4.1.1,1421a. Male. Purchased from an Italian.
N. Mediterranean.
Complete skull and skeleton of an immature animal.
3. 1892. 10.4. 1, 1063d. Presented by C. F. R. Blandy.
Deserta Grande Is. Madeira.
Complete skull and skeleton of an immature animal.
4. 1892. 11. 7. 1, 1063I. Presented by C. F. R. Blandy.
Deserta Grande Is. Madeira.
Complete skull and skeleton of very young animal.
Skin also in collection.
5. 1894.7.27. 1, 1063! Male. Presented by C. F. R. Blandy.
Deserta Grande Is. Madeira.
Complete skull and skeleton of adult animal.
6. 1894.7.27.2, io63g. Female. Presented by C. F. R. Blandy.
Deserta Grande Is. Madeira.
Complete skull and skeleton of adult animal.
Skin also in collection.
7. 1894.7.27.3, 1063I1. Male. Presented by C. F. R. Blandy.
Deserta Grande Is. Madeira.
Complete skull and skeleton of immature animal.
THE MONK SEALS 225
The seal was sent alive from Madeira and lived in the Zoological Gardens
London from 16th July to 28th October, 1894.
Skin also in collection.
8. 1951.4.17.2. No history.
Incomplete skeleton of an immature animal.
No skull.
Skins
1. 1890. 12. 30. 1. Male. Presented by H. C. Hinton and C. J. Cossart.
Bugio, Deserta Grande Is. Madeira.
Skin of an adult animal.
2. 1892. 11. 7. 1, 1063I.
Skin of very young animal noted above.
3. 1894.7.27.2, io63g. Female.
Skin of adult animal noted above.
4. 1894.7.27.3, 1063I1. Male.
Skin of immature animal noted above.
Stuffed animals
1. 1853. 10.6.4, 1063a.
Belonging to type of H. atlantica Gray, noted above.
2. 1 910 . 9 . 27 . 1. Presented by Zoological Society of London.
Presented to Zoological Society by Godfrey Williams.
Madeira.
3. An immature animal with no history.
M. tropicalis
Skull only
1. 1889 .11.5.1, 1064b. Male. Presented by F. D. Godman.
Triangle Is. Gulf of Mexico.
Collected in 1886 by Comision Geografico Exploradoro, Mexico.
Skull and lower jaw of adult animal. Skull complete except for both jugals.
Skin also in collection.
Skull and skeleton
1. 1887 .8.5.1. Collected by H. L. Ward.
Triangle Is. Gulf of Mexico.
Complete skull and skeleton of adult, probably female animal.
2. K.7801, 899c. Presented by F. D. Godman.
Triangle Is. Gulf of Mexico.
Complete skull and skeleton of adult animal. In University Zoological
Museum, Cambridge.
226 THE MONK SEALS
Skin
I. 1889. 1 1. 5. i, 1064b. Male.
Skin of adult animal noted above.
Stuffed animal
1. 1847.2.2.2. Presented by P. H. Gosse.
Jamaica.
Type of Phoca tropicalis Gray, 1850.
M. schauinslandi
i- 32795.
The skull of the type specimen collected on Laysan Id. by Dr. Schauinsland.
The skull is in the Zoological Museum in Berlin and has not been examined,
but photographs have been made available by Dr. K. Zimmermann, and a
very complete set of measurements was published by Matschie (1905).
A stuffed specimen is believed to be in the Bremen Museum.
OSTEOLOGY
A skull belonging to the genus Monachus has the following characters :
1. The skull is broad in proportion to its length.
2. The dorsal surface is convex, sloping backwards and forwards from a point
about halfway along the interorbital region.
3. The interorbital region is broad and parallel-sided.
4. The naso-maxillary region is flattened dorsally.
5. The snout is broad, the sides of the maxillae being almost parallel from the
upper edge of the infra-orbital foramen to the canines.
6. The molars are large and set more or less obliquely, and there are four upper
and four lower incisors.
7. The condyle of the lower jaw is very low, on a level with a line drawn through
the points of the molar teeth.
Comparison of the skulls of M. monachus, M. tropicalis and M. schauinslandi
1. In view of the small number of skulls examined, few general conclusions can be
drawn from the measurements and proportions (Table I), although, as the following
summary shows, skulls of M. monachus tend to be slightly wider in proportion to
their length than skulls of M. tropicalis and M. schauinslandi (Text figs. 5 and 6,
PL 6).
M. monachus. M. tropicalis. M. schauinslandi.
0/ 0/ 0/
/o /o /o
Zygomatic width . . . . 59- 9-70 -3 . 61-7-62-1 . 61-5
Snout width at canines . . 20-9-26-0 . 20-6-20-9 . 20-3
Width at external auditory meatus 53*1-58-5 . 49*8-50-5 . 50 • 5
Width at petrous bones . . 60 • 3-64 -9 . 56 • 3-59 -2 . 59-2
THE MONK SEALS
227
10 CMS.
10 CM5.
Fig. 5. — M. monachus Reg. No. 1894.7.27. 1. a. Dorsal view of skull, b. Ventral view.
Note supernumerary incisors, c. Lateral view of skull and lower jaw.
228
THE MONK SEALS
10 CMS.
10 CMS.
Fig. 6. — M. tropicalis Reg. No. 1887.8.5. i. a. Dorsal view of skull, b. Ventral view,
c. Lateral view of skull and lower jaw.
THE MONK SEALS 229
2. Each nasal of M. monachus ends anteriorly in a V, the point of the V, which
may be slightly rounded, being directed anteriorly, so that the ends of both nasal
bones together form a W. The nasal septum may project slightly beyond the nasal
bones in old specimens. The nasal bones of M. tropicalis are much longer and
narrower and their anterior ends continue the curve made by the premaxillae. The
nasal septum forms a triangular projection. The nasal bones of M. schauinslandi
are as long as those of M. tropicalis, but do not taper to so fine a point posteriorly.
The anterior end of each has the form of an inverted V with the point directed
posteriorly. The nasal septum appears to project a little beyond the end of the
nasal bones (Text-fig. 7).
3. When seen from in front, the lower edge of the infraorbital foramen is wider
than the upper edge in M. monachus, while the reverse is true in M. tropicalis. The
foramen is not visible in the photographs of M. schauinslandi, although Matschie
(1905) says that the lower edge is wider than the upper.
4. There is a well defined tubercle on the maxilla at the anterior margin of the
orbit in M. monachus. This is very small and indistinct in M. tropicalis, and
Matschie (1905) says that M. schauinslandi also has no clear tubercle in this position.
5. In M. monachus the zygomatic branch of the squamosal is not expanded at its
anterior end and lies at an angle of approximately 45 ° with the base of the skull.
The orbital process of the jugal is wide and upwardly directed while the lower branch
is narrower and continues back along the squamosal for some distance.
In M. tropicalis the zygomatic branch of the squamosal is expanded at its anterior
end and is much more nearly vertical. The orbital process of the jugal is very
narrow and curves backwards over the top of the zygomatic branch, while its lower
branch is short and triangular.
In M. schauinslandi the zygomatic branch of the squamosal is similar to that of
M. monachus ; it is not expanded and lies at an approximate angle of 45 °. The
orbital process of the jugal is not quite so wide as that of M. monachus, but is
otherwise similar, and the lower branch is long and backwardly directed (Text-
fig- 7).
6. The posterior end of the palate of M. monachus is U-shaped with a small median
V-shaped incision. In M. tropicalis it is V-shaped, and in M. schauinslandi it forms
a slightly wider V. (Text-fig. 7) .
7. The pterygoid bones of M. monachus are low and ill-defined, and are not visible
when the skull is viewed dorsally. In M. tropicalis they curve widely outwards
and are visible dorsally. In M. schauinslandi they curve out as in M. tropicalis,
but are not visible dorsally.
8. The coronoid process of the lower jaw of M. monachus is wide and concave
internally. That of M. tropicalis is narrow and only very slightly concave. The
coronoid of M. schauinslandi is narrow and like that of M. tropicalis.
9. From the above characters it will be seen that the skull of M. schauinslandi is
more like that of M. tropicalis than M. monachus.
230
THE MONK SEALS
NASAL BONES
C.
ZYGOMATIC ARCH
HIND END PALATE
Fig. 7. — Nasal bones, zygomatic arch and hind end of palate of a. M. monachus, b. M.
tropicalis, c. M. schauinslandi. sq., squamosal ; ju., jugal.
THE MONK SEALS
231
Table I. — Cranial Measurements of Monachus Skulls.
M. monachus.
1951-
4. 17. 1.
1894.
7.27.1.
A
1863
.4.1.1.
a
1894.
7.27.2.
A
1934
•8.5.4-
.a
r
^
(
^
r
^
r
"\
f
mm.
%
mm.
°/
/o
mm.
%
mm.
%
mm.
%
Condylobasal length .
295
100
. 281
IOO
• 273
IOO
. 268
IOO
. 262
IOO
Condylobasilar length
282
95'5
. 270
96-1
. 262
95*9
• 258
96.2
253
96-5
Basal length
279
94-5
• 263
93-5
. 251
91-9
• 251
93*6
241
91-9
Basilar length .
266
90-1
. 252
89-6
. 241
88-2
. 241
89-9
232
88-5
Snout width at canines
7i
24-1
. 68
24-1
• 7i
26-0
• 57
21 -2
■ 55
20-9
Width of skull at front
80
27-1
. 76
27-0
• 85
3i'i
. 60
22-3
64
24-4
end of last upper
molars
Zygomatic width
205
est.
69-4
• 193
68-6
. 185
70-3
. 176
6 5 -6
159
6o-6
Width at upper edge
166
56-2
. 156
55*5
. 150
54'9
• 145
54'i
139
53*1
auditory meatus
Width at petrous
178
60 -3
• 177
62-9
. 165
60 -4
. 167
62-3
. 160
61 • 1
bones
Palatal length .
127
43*1
• ii7
41-6
• 113
4i-3
. no
41 -o
106
40-4
Palatilar length
113
38-3
. 107
38-i
■ 103
37-7
IOO
37'3
97
37-°
Width of occipital con-
65
22-0
• 67
23-8
. 64
23'4
. 65
24-2
■ °5
24-8
dyles
Length of nasal suture
57
19-3
. 46
16-3
• 52
19-0
53
19-7
49
18.7
Length of upper molar
64
21 -6
. 60
21-3
. 64
23'4
57
21 -2
56
21-3
row
M. monachus.
1892.
10.4. I.
1894.
7-27-3-
1853.
10.6.4.
1892.
II .7.1.
1063c.
K
_a_
A
<
A
/
\
*\
r
> k
> »
(
^
mm.
%
mm.
%
mm.
%
mm.
%
mm.
%
Condylobasal length .
260
IOO
. 224
IOO
. 217
IOO
171
IOO
217
IOO
Condylo-basilar length
249
95*7
. 216
96-4
. 212
97-7
164
95*9
208
95 '9
Basal length
241
92-6
207
92-4
. 197
90-8
156
91 -2
197
90-8
Basilar length .
230
88-4
198
88-4
. 191
88-o
M7
85-9
188
86-6
Snout width at canines
61
23'4
52
23-2
. 5i
23*5
44
25'7
50
23-0
Width of skull at front
66
25-3
• 63
28-1
• 63
29-0
56
32-7
67
30'9
end of last upper
molars
Zygomatic width
164
63-1
139
62-1
130
59*9 ■
108
63-2 .
137
63-1
Width at upper edge
144
55*3
126
56-2
123
56-7 •
IOO
58-5 •
122
56-2
auditory meatus
Width at petrous
163
62-6
138
6i-6
136
62-7 .
in
64-9 .
136
62-7
bones
Palatal length .
102
39'2
90
40*1
91
41-9 .
73
42-7 .
92
42-4
Palatilar length
92
25-3
82
36-6
85
39*2 .
65
38-0 .
84
38-7
Width of occipital con-
69
26-5
63
28-1
. 62
28-6 .
54
31-6 .
57
26-3
dyles
Length of nasal suture
48
18-4
53
23-6
—
— .
38
22-2 .
39
17-9
Length of upper molar
56
21'5
5i
22-7
• 52
23*9 •
ca^6
26-9 .
55
25-3
row
2 32
THE MONK SEALS
Table I — cont.
M. tropicalis.
M.
889c.
schauinslandi.
(Cambridge)
1887
8.5.1.
1889.
II.5-I-
32795.
A
._A
-* ^
>
r
\
(~<
(
r
^\
mm.
%
mm.
%
mm.
%
mm.
%
Condylobasal length
272
100
269
IOO
267
IOO
265
IOO
Condylo-basilar length .
263
96-7
262
97*3
258
96-6 .
255
96-2
Basal length
260
95*6
254
94.4
250
93'6
252
95'i
Basilar length
250
91-9
247
91-8
241
90-2
241
90-1
Snout width at canines
56
20-6
56
20-8
56
20-9
54
20-3
Width of skull at front end o
f 77
28.3
74
27'5
80
29-9
70
26-4
last upper molars
Zygomatic width .
168
6i-8
167
62-1
165
61 -7
163
61.5
Width at upper edge auditors
T I36
50-0
136
50-5
133
49-8
134
50-5
meatus
Width at petrous bones
153
56-3
154
57'2
■ 151
56-5
157
59'2
Palatal length
III
40-8
116
43-i
112
41-9
122
46-0
Palatilar length .
I02
37'5
no
40-8
104
38-9
112
42-2
Width of occipital condyles
■ 67
24-6
69
25-6
63
23*5
66
24*9
Length of nasal suture .
• 67
24-6
65
24*1
61
22-8
61
23-0
Length of upper molar row
66
24-3
• 65
24-1
66
24.7
M. monachus
T
eeth
Dental formula : . 2
1 5
1-, c
2
i' m
Upper. — The incisors are large and pointed, the point directed posteriorly. The
outer incisors are larger than the inner and there is a well developed cingulum on the
inner surface of all four teeth.
The canines are of moderate size with a slight ridge down the posterior surface.
The molars are large, the 3rd being the largest and the others decreasing in size in
the order 3, 2, 4, 1, 5, except in the largest skull (1951.4.17.1) where the 5th is
slightly greater than the first, and the others decrease in size from before backwards.
Although the molars are set obliquely they do not overlap one another. The first
molar makes an angle of 20 with the median palatal suture, the 2nd molar 40 , the
3rd molar 6o°, the 4th molar 70 and the 5th molar no°, so that its palatal surface
is facing posteriorly. The external edge of the palate forms an angle of 20 with
the median palatal suture. These measurements are from 1892.10.4.1, and are
compared with a Phoca vitulina 1919.7.7.3260, where the teeth are set in line at an
angle of 20 . There is a slight obliquity in the teeth of some of the other P. vitulina
skulls, but this is not nearly so pronounced as in M. monachus. All the molars are
double rooted except the first. The molars have a large central cusp and single
THE MONK SEALS 233
smaller anterior and posterior cusps. The small cusps on the 5th molar are less
distinct and the posterior one may be lacking. There is a well developed cingulum
on the inner surface.
Variation. — In the nine specimens with fully erupted teeth, variations in the
number of upper incisors occur in five of them.
1. 1853. 10. 6. 4, 1063a.
There is a small extra tooth just posterior to the first left incisor.
2. 1063b.
Although the tooth is missing in the specimen, the alveolus shows evidence of an
extra tooth posterior to the first right incisor.
3. 1894.7.27. 1, 10631.
On the right side there is a small incisor median to the first and in the same line
with it. The first incisor is the same size as that on the left side, though
farther from the mid line. The second incisor on the right side is separated
from the first by a gap, and leans away from it, towards the canine. It is
considerably smaller than the left second incisor. Posteriorly and internally
to the first and second right incisors is part of the root of a tooth larger than
the left second incisor and smaller than the canine. The crown has been
broken off since the animal died.
4. 1934.8.5.4.
On the left side a small incisor is set directly behind the first.
5. 1951.4.17.1.
On the left side a small incisor is situated just posteriorly to the first.
Disease. — Only one specimen shows any sign of disease in the teeth. 1863 .4.1.1,
1421a — the second right incisor has lost most of the crown, the tooth is hollow, and
there is some lumpy growth on its anterior surface. The parts of the premaxilla
round the tooth have fallen away.
Lower. — The first incisors are considerably smaller than the second, they are set
internally and posteriorly to them and are in a more or less recumbent position.
They are similar to the upper incisors in shape ; the canines are also similar.
The third molar is the largest, the others decreasing in size in the order 3, 2, 4, 5, 1,
so that the 1st molar is the smallest. The molars are set obliquely and do not over-
lap one another. The first molar is set directly behind the canine, so close that the
two alveoli are confluent, but the molar is not on the inner surface of the canine as
in the upper jaw. The angle the teeth make with the symphysial line is not so
varied as in the upper jaw. The 1st molar makes an angle of 30 , the 2nd 40 , the
3rd 40 , the 4th 6o°, and the 5th 20 . As the ramus itself is at an angle of 20 the
5th molar is in line with the jaw. These measurements are from 1863 .4.1.1, 1421a.
All the molars are double rooted except the first. The shape of the lower molars is
similar to that of the upper, except that the posterior cusp on the 5th molar is
generally present.
There are no variations in number, and no disease is present. The surface of all
the teeth, both upper and lower, is slightly rugose.
ZOOL. 3, 5. 17
234 THE MONK SEALS
M. tropicalis
Dental formula : . 2 1 5
l i- c ? m 5
Upper. — The incisors are set in a straight line across the front of the premaxillae.
They are similar in general shape to those of M. monachus, although there is more of
a " waist " at the junction of root and crown. The canines are similar to those
of M. monachus.
The molars are large, the 3rd is largest and the others decrease in size in the order
3, 2, 4, 1, 5. They are hardly oblique, the 2nd molar being the most so. The molars
are double rooted except the first, and the last also appears to have only one root in
the two British Museum specimens. The molars have a low central cusp, and one
anterior and two posterior smaller cusps. The 5th molar has only a single posterior
cusp. All the teeth have a well developed cingulum and the crown is slightly rugose.
Variation and disease. — In the three skulls examined there is no variation or disease
in upper or lower teeth.
Lower. — The first lower incisors are smaller than the second, are set internally and
posteriorly to them and are in a recumbent position. In shape they are similar to
the upper incisors. The lower canines are similar to the upper ones.
The 3rd molar is the largest and the others decrease in size in the order 3, 4, 2, 5, 1,
although molars 3, 4 and 2 are very much of the same size. The teeth are set in the
line of the jaw, not obliquely. All the molars are double rooted except the first.
The lower molars are similar in shape to the upper except that the cusps on the 1st
are rather indistinct, and there is only one small posterior cusp on the 5th.
The teeth of M. tropicalis examined seem to be more worn than the teeth of
M. monachus.
M. schauinslandi
Dental formula : . 2 1 5
1 -, c -, m ^
2 i' 5
Upper. — The incisors are set in a straight line across the front of the premaxillae,
and as far as can be seen from the photographs, the setting and shape of the teeth are
similar to those of M. tropicalis. The molars are not set obliquely and seem to have
a main low cusp, a single small anterior cusp and two small posterior cusps.
Lower. — The lower teeth appear to be similar to those of M. tropicalis.
Summary of the differences between the teeth
1. The incisors have a very pronounced " waist " at the junction of root and
crown in M. tropicalis and M. schauinslandi.
2. The molars are set very obliquely in M. monachus.
3. The molars of M. monachus have a large central cusp and single smaller
anterior and posterior cusps. Those of M. tropicalis and M. schauinslandi have a
low central cusp, a single small anterior cusp and two small posterior cusps.
THE MONK SEALS
235
4. The incisors are set straight across the premaxillae in M. tropicalis and M.
schauinslandi , but on a slight curve in M. monachus.
Teeth of a very young M. monachus
There is no exact information about the skull 1892. n. 7.1, 1063I, but from its
size and the condition of its teeth it is probably new-born. X-rays of the teeth
were taken (Nos. 764 and 765) (Text-fig. 8).
5 CMS.
Fig. 8. — Tracings from X-ray photographs of the teeth of a young M. monachus, Reg. No.
1892. 1 1. 7. 1. a. Ventral view. b. Oblique view. c. Lower jaw.
236 THE MONK SEALS
Milk teeth
These are seen more clearly in the skull itself than in the X-ray. Six milk teeth
are visible in the upper jaw, three on each side. They are very small, the largest
being 2 mm. long ; they have no roots and are present in the dried gum over the
unerupted permanent teeth. The milk molars are placed behind their permanent
successors. In the upper jaw milk molars 3 and 4 are present on the right side, and
2 and 3 on the left side. The two outer milk incisors are present. In the lower jaw
the only milk tooth present is mm 2 on the right side.
There is no milk predecessor for the first molar in either upper or lower jaws.
Milk dentition : . 2 1 3
1 -, c -, m -
2' i 1 3
Permanent teeth
The tips of all the permanent teeth are through the layer of dried skin on the
skull, though probably apart from the upper canines, none of them would have
pierced the gum in the living animal.
The permanent molars are large, but are normal in shape and position. This
specimen shows an interesting variation in the number of incisors. The upper jaw
has seven incisors, three of which are supernumerary. On the left side the outer
incisor is in its normal position, the inner incisor is pushed a little further away from
the mid line than is normal, and on its median surface, slightly posteriorly is a small
extra tooth. On the right side the outer incisor is displaced posteriorly by an extra
tooth which is between it and the canine, and between the inner incisor and the mid
line is another extra tooth of the same size as the small extra one on the left side.
The larger extra tooth on the right has the same size and appearance as the inner
incisor.
The lower permanent teeth are normal.
Supernumerary bones in the skull.
Three of the youngest skulls of M. monachus (1892 .11.7.1, 1063c and 1894 . 7 . 27 . 3)
have supernumerary bones in the cranium. The youngest specimen (1892. n. 7.1)
has two bones symmetrically placed in the back of the skull, each bone bounded by
exoccipital, parietal and supraoccipital. These bones are frequently found in the
young of other seals and have been known as the tabulare, on the assumption that
they were homologous with the tabulare of the reptilian skull. Doutt (1942)
mentions their presence as well as that of two small adjacent bones, in skulls of Phoca
vitulina and P. hispida and after quoting various authors who have considered the
relationships of these bones with those in the reptilian skull, he thinks " that it is
better, for the present at least, to consider these extra bones in the occipital region
of the seal as being of the nature of fontanelle bones rather than to try to homo-
logize them with elements in the reptilian skull ". He says that these " extra-
occipital " bones cannot be considered as Wormian bones for they are too
symmetrical and too regularly situated.
THE MONK SEALS
237
Undoubted Wormian bones are also found in this skull and in the two other young
ones mentioned. In 1892. n. 7.1 they take the form of three small bones, two
posteriorly and one anteriorly, at the junction of the two parietal bones with the
frontal bones. In 1063c there is a single triangular bone in the same position.
Also in the same position in 1894.7.27.3 there are two narrow bones one behind
the other (Text-fig. 9).
POSTERIOR
ANTERIOR
B.
Fig. 9. — a. "Extra-occipital" bones in M. monachus Reg. No. 1892. 11. 7.1. b.
Wormian bones in M. monachus Reg. No. 1892. 11. 7.1 ; c. Wormian bones in M.
monachus Reg. No. 1063c ; d. Wormian bones in M. monachus Reg. No. 1894.7.27.3.
p., Parietal ; /., frontal ; ex., exoccipital ; soc, supraoccipital.
Scapula (Text-fig. 10)
The scapula of Monachus is triangular in shape, the extreme antero-posterior
length being greater than the height. In this character it differs from the scapulae
of other Phocids where the anterior edge is much longer and, especially in the
southern Phocids, much more square in shape. The junction of the posterior and
dorsal edges is not hook-shaped. The spine is low and poorly developed like those
238
THE MONK SEALS
of Mirounga and the southern Phocids, and very unlike the plate like spines of
P. vitulina and H. grypus. The acromion process is well developed and resembles
that of the southern Phocids, and the glenoid cavity is narrow and kidney-shaped.
The outer surface of the scapula is convex anteriorly to the spine and concave
posteriorly to it. Scapulae of P. vitulina and H. grypus are convex posteriorly
and concave anteriorly.
10 CMS.
Fig. io. — Scapulae of a. M. monachus Reg. No. 1894.7.27. 1.
b. M. tropicalis 1887.8.5. 1.
In young M . monachus and in M. tropicalis the anterior edge of the scapula sweeps
round in a continuous curve from the neck, forwards and then back along the dorsal
edge. In adult M. monachus the anterior edge is more squared off. The scapula
of M . tropicalis is more elongated antero-posteriorly than that of M. monachus.
The separate origin of the coracoid is visible in three of the youngest specimens of
M. monachus. In the youngest scapula (1892. 11. 7.1) it does not appear to have
THE MONK SEALS 239
begun to ossify. Scapulae 1894.7.27.3, and 1063c have the cartilage still present
over the glenoid cavity and the coracoid is visible as a distinct bone embedded in it.
Scapula 1892. 10.4. 1 has no cartilage, the coracoid has fused to the anterior edge of
the glenoid cavity, but is still recognizable as a separate entity.
Humerus
The humerus is short and robust. In common with all the southern Phocids
there is no supracondylar foramen. This foramen is present in Phoca, H. grypus,
E. barbatus and C. cristatus. The deltoid ridge is strongly developed and extends
nearly the whole length of the shaft. The deltoid rugosity on the external surface
of the ridge is more prominent in M. monachus than in M. tropicalis, but does not
overhang to so great an extent as in Phoca, Halichoerus and Erignathus. The
bicipital groove is broad and shallow. In Monachus and the southern Phocids
there is only a very poorly marked supinator ridge, while this is well developed in
the northern Phocids. For bones of approximately the same length, the humerus of
M. tropicalis (PI. 7,a) is slightly more slender than that of M. monachus. The
distal articular surface is narrower and appears to be slightly more oblique than in
M. monachus. In M. tropicalis, but not in M. monachus there is a very slight
anconeal fossa.
Radius and Ulna.
The radius is laterally flattened and bowed forward, so that its hinder edge is
concave. The ulna is compressed laterally, with an expanded olecranon process
and a concave hinder margin. The ulna of M. monachus is slight y more stoutly
built than that of M. tropicalis, but presents no special modifications.
Manus (PL 7,a)
The carpus consists of seven bones : scapholunar, trapezium, trapezioid, os
magnus, unciform, cuneiform and pisiform. The length of the digits, and of the
metacarpals decreases in size from the first to the fifth. The first digit is strongly
built, the fifth slightly less so, and the second, third and fourth of equal thickness.
The terminal phalanges are grooved for the insertion of claws. An X-ray taken of
the manus of an immature M. monachus (1894 . 7 . 27 . 3) shows the presence of distal
epiphyses on all five metacarpals, but a proximal epiphysis on only the first. All
the phalanges except the terminal ones have epiphyses on both ends, and a proximal
epiphysis is visible only on the first terminal phalange, it not yet having ossified
on the remaining phalanges.
Pelvis and sacrum
The innominate bones are of the typical Phocid pattern with short recurved ilia
and long ischia and pubes. The symphysial area is very small and appears to be
mainly cartilaginous. The innominate bones of M . monachus have thick flattened
ilia which are not excavated on their exterior surfaces as in Phoca and Halichoerus,
240
THE MONK SEALS
When compared with the innominate bones of other Phocids it is seen that the ilia
of M. monachus are everted and also are rotated upwards relative to the plane of
the pubes and ischia, so that the external surfaces of the ilia face slightly posteriorly
and dorsally. In Phoca and Halichoerus and also in the southern Phocids the ilia
are everted in approximately the same plane as the pubes and ischia. The ischium
of M . monachus is strongly built, approximately circular in cross section until about
the level of the ischiatic spine, when it continues as a flat bar. The pubis is also
thicker at its origin and flattens out posteriorly to meet the ischium. The ischium
forms the upper and practically the whole of the posterior border of the obturator
foramen before it meets the pubis.
A comparision of the innominate bones of Phocids shows that in the northern
genera Phoca, Halichoerus and Erignathus the obturator foramen is long and narrow
and the width of the innominate at the level of the ischiatic spine less ; in the
southern genera Ommatophoca, Hydrurga, Leptonychotes, Lobodon and Mirounga the
obturator foramen is broad and the width of the innominate correspondingly
greater. The proportions of the obturator foramen given in the table below indicate
a closer relationship of Monachus with the Southern than the Northern Phocids
(PL 8).
Measurements of the Obturator Foramen
Extreme
anteroposterior
Greatest
Proportion
length.
width.
of width
(mm.)
(mm.)
to length.
P. vitulina .... 96
3 1
32-3
H. grypus
100
26
26-0
E. barbatus .
112
37
33-o
L. weddelli
103
43
41-7
H. leptonyx .
107
57
53*3
L. carcinophagus
97
53
54*6
0. rossi
88
45
5i-i
M. monachus
io63g 94
5i
54'3
io63f 98
55
56-1
1421a 82
50
60 9
M. tropicalis
1887.8.5. 1 ... 112
50
44-6
899c.
114
52
45-6
In M . monachus the pectineal tubercle and the ischiatic spine are well developed,
the acetabulum is deep and circular with a well marked cotyloid notch. The
innominate bones articulate with two sacral vertebrae, but in the three adult M.
monachus, two have four vertebrae fused to form the sacrum, and one — the
youngest — has three. The wings of the sacrum are broad and strong, and
articulate on their posterior surfaces with the ilia.
THE MONK SEALS 241
In most respects the pelvis of M. tropicalis is similar to that of M. monachus, but
the size of the ischium and pubis differs considerably. The pubis is extremely
strongly built, is circular in cross section, tapers slightly towards its posterior end
and does not flatten out except at the symphysial area. The ischium is much more
slender than the pubis, and posterior to the ischiatic spine it is only a very narrow
bar, triangular in cross section.
Femur
The femur is short and flattened antero-posteriorly. The greater trochanter is
large and thickened and is separated from the head by a groove which is deeper
and more distinct in M. monachus than in M. tropicalis. In common with all Phocids
except Phoca and Halichoerus there is no trace of a digital fossa. The distal end of
the femur is very broad because of the great development of the epicondyles. In
general shape, and in particular in the great width in proportion to its length, the
femur of Monachus is more like that of the Southern than the Northern Phocids.
Patella
The patella of M. monachus is a small flat bone articulating with the femur. The
measurements of specimen i033f are : Greatest antero-posterior length 26 mm.
Greatest height anteriorly 15 mm. The patella of M. tropicalis, while of the same
general length, is, particularly in the Cambridge specimen, higher and more
pyramidal. The measurements of the British Museum and Cambridge specimens
respectively are : Greatest antero-posterior length 30 mm., 24 mm. Greatest
height anteriorly 20 mm., 28 mm.
Tibia and Fibula
The tibia and fibula are of the usual pinniped form, the two bones being fused at
the proximal end, but separate from each other at the distal end. In only one of
the M . monachus in the collection (io63g) is one of the fibulae fused to the tibia, so
this fusion must be one of the last to occur. The posterior tibial fossa is less concave
than in any of the other Phocids examined, but the tibial spine and the anterior
tibial fossa are quite well marked. Both the tibia and the fibula articulate at their
lower ends with the astragalus.
Pes (PI. 7,b)
The tarsus consists of seven bones : astragalus, calcaneum, navicular, cuboid and
external, middle and internal cuneiform bones. The length of the digits and of
the metacarpals decrease in size in the order 1, 5, 2, 4, 3, the middle digit being the
shortest. The outer digits are the most strongly built. The terminal phalanges
are grooved for the insertion of small claws. An X-ray taken of the pes of a young
M. monachus (1894.7.27.3) shows the presence of distal epiphyses on all the
metatarsals, and a proximal one on the first only. Epiphyses are visible on both
242
THE MONK SEALS
ends of all the phalanges except the terminal ones. Epiphyses have not yet ossified
on the proximal ends of any of the terminal phalanges except the first.
Vertebral Column
The vertebral formula is as follows :
M . monachus
Cervical.
Thoracic.
Lumbar.
Sacral.
Caudal
1892. 1 1. 7. i, 1063I.
7
15
5
2
3 +
1894.7.23.3, 1063I1. .
7
15
5
3
11
1894.7.27.2, io63g.
7
15
5
4
11
1894.7.27. 1, io63f
7
15
5
4
11
1892. 10.4. 1, 1063d
7
15
5
3
11
1863. 4. 1. 1, 1421a
7
15
5
3
11
1063c
7
15
5
2
10
M. tropicalis
1887.8.5. 1
7
15
5
3
12
899c .
7
15
5
3
12
Cervical vertebrae
M. monachus. — The centra are approximately circular in cross section and the
ventral surfaces have a median keel in all except the atlas. The neural arches are
narrow antero-posteriorly, the widest (the 7th) being 20 mm., and the neural spines
increase from a hardly perceptible point on the third vertebra to a spine 38 mm.
high on the seventh. The transverse processes are perforated by the vertebrarterial
canal in all except the seventh. The transverse processes of cervical vertebrae 3-6
inclusive are, in all the Phocidae examined, divided into two branches, a dorsal
transverse element which is more or less at right angles to the median plane, and
directed slightly posteriorly, and a ventral costal element which is directed more or
less vertically downwards and is expanded antero-posteriorly into a plate which is
greatest in the sixth vertebra. In M . monachus the transverse processes leave the
centrum at an angle of approximately 45 ° and are not divided into two branches
(Text-fig. 11).
The cervical vertebrae of M. tropicalis, while conforming to the same general
pattern as those of M. monachus, are different in several minor respects. They
give the general impression of being more finely built than those of M. monachus.
The transverse process of the atlas is not so massive and the vertebrarterial canal is
much larger, the neural spine of the axis is not so high and does not project so far
backwards. The transverse process of the axis is thin and pointed, those of verte-
brae 3-6 are divided into two branches, the costal element not being so expanded
as in Phoca and directed not vertically, but laterally at an angle of approximately
45°, and inclining posteriorly. The general shape of the cervical vertebrae and in
particular that of the transverse process is more like Leptonychotes than Phoca.
It is interesting to note that both in the British Museum and the Cambridge
specimens of M . tropicalis the neural arches of the third and fourth vertebrae do
THE MONK SEALS
243
not meet dorsally. In the British Museum specimen the two sides of the neural
arch of the fifth vertebra are fused, but the lateral tips of the spine are curved
outwards, to give a bifurcated tip to the spine. The lateral tips of the sixth spine
are less curved. In the Cambridge specimen the two sides of the fifth neural arch
Fig. 11. — a. 5th cervical vertebra of M. monachus Reg. No. 1894.7.27.2. b. 5th
cervical vertebra of M. tropicalis Reg. No. 1887.8.5. 1. c. 2nd lumbar vertebra of
M. monachus Reg. No. 1894.7.27.2. d. 2nd lumbar vertebra of M. tropicalis Reg.
No. 1887.8.5. 1.
meet, but are not fused, and the tips curve outwards. The spine of the sixth is
fused with curved tips, and the spine of the seventh in both animals is normal. A
similar non-fusion of the neural arch occurs in M. monachus 1063c, where the two
sides of the third neural arch fail to meet. The remaining arches are normal and
do not have the bifurcated appearance of those of M. tropicalis. Ward (1887b)
244
THE MONK SEALS
says, describing M. tropicalis, "At no time does this seal raise its head as much
above the line of its back as does the harbor seal : the flexibility of its cervical
vertebrae appearing to be quite restricted ". It is difficult to know whether this
is a character that applies to M. tropicalis in general, or only to the animals that
Ward saw ; or whether it has any connection with the non-fusion of the neural
spines, as the two animals in which this character is present were both collected by
Ward from a single group of animals.
Thoracic vertebrae
The thoracic vertebrae of M. monachus articulate with fifteen pairs of ribs. Ribs
i and 2, and n, 12, 13, 14 and 15 articulate only with the corresponding thoracic
vertebra, but the capitular articulation of ribs 3-10 inclusive overlaps on to the
centrum of the vertebra in front. Ribs 1-13 inclusive articulate by both capitular
and tubercular heads, the two articulations getting closer together until ribs 14 and
15 articulate by means of a single head only. The transverse processes are
prominent, but diminish in size from the first to the tenth vertebra and are hardly
visible from the eleventh to the fifteenth. The neural spines are high and pointed
in the first five vertebrae, but get progressively smaller and more backwardly
inclined.
Between thoracic vertebrae 9 and 10 in specimen 1863 .4. 1 . 1, the anterior common
ligament has ossified in the form of a bony plate, which is fused to the centrum of
the tenth vertebra and extends anteriorly for 4cm. beyond it. Evidences of a
similar abnormality, though to a much lesser extent, are present on the ventral
surfaces of the centra of vertebrae n, 12 and 13.
The thoracic vertebrae of M. tropicalis are practically the same as those of M.
monachus except that the neural arches are wider and the neural spines are shorter
and less pointed.
Measurements of thoracic vertebrae
Length medially of neural arch of 10th vertebra
Least width ,, ,,
Height* of neural spine of 1st vertebra
5th
10th
Length of centrum of 1st „
„ „ „ „ 10th
M. monachus. M. tropicalis.
io63g.
1887.8.5. 1.
mm.
mm.
48
46
25
34
58
5i
65
52
54
46
41
40
52
5i
Height not taken vertically, but medially along spine.
Lumbar vertebrae
The lumbar vertebrae of M. monachus have large heavy centra that are slightly
concave ventrally. The transverse processes are prominent and project anteriorly
and ventrally. The cephalic articular processes are large and are directed obliquely
THE MONK SEALS
245
anteriorly, but the caudal articular processes are extremely small and thin, and may
not even reach the vertebra behind. The neural spines are stoutly built and are
directed slightly posteriorly.
The right transverse process of the first lumbar vertebra of M. monachus io63f
is normal, but the left is stout, truncated, and its distal end forms an articulation
for a small triangular " pleurapophysial " ossicle 34 mm. in length.
A condition of chronic osteo-arthritis is present on the posterior half of the centrum
of the fifth lumbar vertebra of M. monachus io63g, and this condition has spread to
the centrum of the first sacral vertebra.
The lumbar vertebrae of M. tropicalis are in general like those of M. monachus,
though the articulations are normal, the caudal articular process overlapping to a
considerable extent the cephalic articular surface of the next posterior vertebra.
The neural spines are laterally flattened, low, rounded, and either vertical or
inclined slightly anteriorly (Text-fig. 11).
Caudal vertebrae
Except for the first two or three, the caudal vertebrae are without a neural arch,
and both the number and size of the processes, and also the size of the centrum,
diminish in size from before backwards.
Ribs
The articulations of the ribs with the thoracic vertebrae have already been
discussed. In M. monachus (io63f) the ribs increase in length from the 1st (yy mm.)
to the nth (263 mm.) and then decrease to the 15th (209 mm.). In M. tropicalis
(1887. 8. 5.1) similar measurements are 1st (71 mm.), nth (255 mm.) and 15th
(189 mm.).
The adult specimens of M. monachus in the collection lack the cartilaginous
portions of the ribs. An immature M. monachus (1063c) has cartilaginous ribs
attached directly to the sternum from bony ribs 1-9 inclusive. Cartilaginous ribs
10-15 inclusive turn forward and lie against the cartilaginous portion of the rib
in front.
Cartilaginous ribs 1-10 inclusive of M. tropicalis (1887. 8. 5.1) are attached
directly to the sternum, 11 and 12 are long and lie against the cartilaginous portion
of the ribs in front, but the cartilaginous portions of ribs 13-15 are very short,
between 35 and 70 mm. long, and are unattached ventrally.
The first cartilaginous rib articulates with the manubrium of the sternum, the
remaining nine with the cartilages between the sternebrae, ribs both 8 and 9
articulating with the last cartilage.
Sternum
The sternum of both M. monachus and M. tropicalis consists of nine sternebrae,
although there are only eight present in M. monachus 1063c. The sternebrae are
dorso-ventrally flattened and more or less quadrate in shape, the first and last being
246
THE MONK SEALS
more elongated. The xiphisternum is prolonged posteriorly into two cartilaginous
extensions, the ends of which are expanded and joined together posteriorly.
GROWTH
In view of the small number of specimens available no attempt at age determina-
tion has been made but it has been possible to do a certain amount on the growth
of the skull and skeleton.
Following Doutt's (1942) method for estimating the " suture age " of the skull
the following table was drawn up. (Table II). As suture closure is a gradual
process the degree of closure has been given a value : 1 for open, 2 for less than
half closed, 3 for more than half closed, and 4 for completely closed, and the total
for each skull is known as the " suture age " for that specimen.
Table II. — Suture Ages of the Skulls
T3
12
a
'0
'0
a
a
■a
CD
i
6
CD
U
CD
co
'So
73
+->
CD
I
&
1
[Oi
'O
O
O
*c3
4->
1
6
co
O
1
W
|
f
oc5
O
O
O
CO
1
,9
6
'0
1
CO
cd
!S
'0
CD
■a
CO
1
73
O
a
h- 1
6
CD
1
|
6
a
2
to
3
8
1
•a
bo
3
•— >
■a
¥
CO
O
g-
>>
PS
1
a
i
1
CD
9
9
in
3
1
6
>>
•d
d
6
M. monachus :
1892. 11. 7. 1
I
I
I
1
I
1
1
12
. 171
1063c
I
I
I
1
I
1
1
12
. 217
1894.7.27.3 <J
2
I
I
1
I
1
1
13
. 224
1892. 10.4. 1
4
I
2
1
I
1
1
16
. 260
1934.8.5.4
4
2
2
2
I
1
1
18
. 262
1894.7.27.2 $
4
4
4
4
4
4
I
1
1
30
. 268
1863. 4. 1. 1 £
4
4
4
3
4
1
I
1
1
26
• 273
1894.7.27. 1 <J
4
4
4
4
4
2
I
1
1
28
. 281
1951.4.17.1
4
4
4
4
4
4
I
1
1
29 +
• 295
M. tropicalis :
1889. 11. 5. 1
4
4
4
2
4
1
I
1
1
I
I
1
25
. 267
1887.8.5. 1
The ore
4
.er of
4
sutur
4
3 CIOSI
3
are is
4
as f ol
4
lows :
I
3
1
• 3
I
1
33
. 269
1.
Exocc
dpita
— bas
ioccip
>ital.
2.
Occip
ito — ]
Dariet
il.
3-
Squar
noso-
-paric
stal
Sagittal.
Coronal.
Basioccipito — basisphenoid.
7. Basisphenoid — presphenoid.
In Table II the skulls are placed in order of condylo-basal length, and it can be
seen that the one known female skull of M. monachus has a higher " suture age " in
THE MONK SEALS
247
proportion to its condylo-basal length than the other skulls, i.e. it matures at a
smaller size. For this reason it seems likely that the skull and skeleton of M.
tropicalis 1887.8.5. 1 are also from a female.
Table II also shows that the series of skulls of M. monachus, although more or
less uniformly distributed over a size range, is divisible by " suture age " into two
discontinuous groups — those with " suture ages " 12-18 and those from 26-30.
This is clearly shown in the table of percentage increases below, from which it is
also evident that while the " suture age " increases from 12-18, an increase of six
units, there is a general increase in the size of all the components of the skull, the
average amount being 53% or 9% per unit of " age ". During the period when the
" suture age " increases by four units from 26 to 30 there is also an increase in all
the components with one exception, but the average rate of increase per unit of
" age " is now only 4.5%. A diminution in the rate of increase with age is to be
expected, but one component — the cranium — appears virtually to have ceased
growing by the time a " suture age " of 18 is reached ; there are no crania in the
26-30 group larger than the largest individuals in the 12-18 group.
Width
at
Condylo-
Percentage increases.
external
basal
Inter-
Zygo-
audi-
Suture
length,
Cranium
orbital
Snout Width at
matic
tory
age."
mm.
length.
length.
length. canines.
width.
meatus.
12-18
171-262
46
77
• 59 • 39
52
. 44
19-25
—
. Negative
13
8 . Negative .
7
0-7
26-30
268-295
2
11
. 24 . 25
17
• 15
The most striking feature of the above table is that, despite the absence of any
large size discontinuity in the series, there is a complete absence of any specimen in
the large " suture " group 19-25 ; the largest specimens of the " younger " 12-18
group are almost as large as, and in some components larger than the smallest
individuals of the " older " 26-30 group.
It is not likely that growth proceeds in this erratic fashion and therefore it is
concluded that the " suture age " is not a rectilinear age index. It seems clear
that at a certain stage of growth there is a very rapid suture closure. After this
stage is past the skull components continue growing with the notable exception of
the cranium. Skulls with a condylo-basal length of less than ± 265 mm. have a
rapidly growing cranium and during this growth-phase the occipito-parietal,
sagittal, coronal and basioccipito-basisphenoid sutures are closing. At ± 265 mm.
condylo-basal length there is rapid suture closure affecting the occipito-parietal,
squamoso-parietal, sagittal, coronal, basioccipito-basisphenoid and basisphenoid-
presphenoid sutures, and cranial growth ceases. After this, growth of the facial
elements continues and the interfrontal, intermaxillary, fronto-maxilla, jugal-
maxilla, squamosal- jugal and palato-maxilla sutures are open in all the skulls of
M. monachus in the collection.
It is possible that the " suture age " figures give an approximation to a rectilinear
age index if the 19-25 period is eliminated, and if this is done the relative " ages "
248
THE MONK SEALS
of the skulls are 12 : 12 : 13 : 16 : 18 : 23 : 19 : 21 : 23. The " suture age " 12
includes two skulls which are both young but which are clearly at very different
stages of growth. The youngest skull — 1892. n. 7.1 — has not been included in the
graph as it is so much younger than its " suture age " indicates, this criterion being
invalid for very young (and probably very old) skulls.
Plotting the skull measurements against this time scale (Text-fig. 12) gives a
Condylobasal
Ungth
Zygomatic
width
Auditory
width
IR 13 14 15 Ifc 17 18 19 20 21 XX Z7>
Suture age
Fig. 12. — Graph showing relative grtfwth rates of different regions of the skull.
THE MONK SEALS
249
picture which is not an unreasonable one, and one which suggests certain sex
differences. Even if the time scale is only an approximation, plotting all the bones
to this scale gives a series of curves that are directly comparable. The slope and
shape of these curves show the relative growth rates and growth patterns of the
different elements.
From a visual inspection of a series of skulls of M. monachus of ascending size, it
is obvious that the various regions of the skull do not all grow at the same rate.
The cranium for instance, while it increases in length, does not do so nearly so quickly
as the facial region. Measurements of the cranial, interorbital and snout regions
were taken between uprights and these, together with width measurements are seen
in Table III. The interorbital region is measured to the maxillary tubercle at the
anterior edge of the orbit.
The logarithmic values of the various measurements were plotted in turn against
the logarithmic value of the condylo-basal length. The points fall approximately
about a straight line, showing that there is allometric growth. This being so, it
may be taken that the rates of growth of the various parts of the skull relative to
the condylo-basal length satisfy the equation y=bx fc , where x is the condylo-basal
length, y the length of the part, b the fractional coefficient (the value of y when
x=i) and k the growth coefficient.
Registered No.
M. monachus
1892. 11 .7.1
1063c
7-27-3
10.4. 1
8.5.4
7.27.2
4. 1.1
7.27.1
4-17-1
1894
1892
1934
1894
1863
1894
1951
Table III
Condylo- Inter-
basal Cranium orbital Snout
length. length. length. length,
mm. mm. mm. mm.
171 . 85
217 . 104
224 . 101
260 . 124
262 . 118
268 . 112
273 . 114
281 . 113
295 . 117
Snout
width at
canines.
mm.
Zygo-
matic
width,
mm.
. 40
46
44
108
. 58
55
50
137 •
61
62
52
139
68
68
61
164
7i
73
55
159 •
81
75
57
176
80
79
7i
185 •
. 89
79
68
193
. 87
9i
7i
205
Width at
upper edge
external
auditory
meatus,
mm.
100
122
126
144
139
145
150
156
166
The growth coefficient is the tangent of the angle between the horizontal and the
line joining the points. Values of k greater than unity indicate an increasing rate
of relative growth, and those less than unity the converse. The following are the
values of k for the various parts :
Cranium length . . . .0-62
Interorbital length
. i-8i
Snout length
. 1-32
Snout width
• 0-95
Zygomatic width .
• I'I5
Auditory meatus width
. 0-85
ZOOL. 3, 5.
18
250 THE MONK SEALS
These figures bear out the visual evidence as they show that the interorbital and
snout regions, and the zygomatic width have a much faster relative growth than
the rest of the skull, while the cranium, the snout width and the width of the cranium
between the external auditory meatuses grow at a much slower rate.
Although fusion of the bones of the skull commences at an early age, the epiphyses
of the skeleton do not fuse until comparatively late. The following is the order of
fusion of the epiphyses and skeletal elements :
i. Pelvic elements.
2. Sacral vertebrae.
Humerus (distal).
Femur (proximal).
5. Vertebral epiphyses.
Humerus (proximal).
Radius (proximal).
Femur (distal).
Tibia (proximal).
Fibula (proximal).
11. Ulna (proximal).
Metacarpal epiphyses.
Metatarsal epiphyses.
14. Fibula to tibia.
15. Radius (distal). "^
Ulna (distal). I Not fused in any specimen
Tibia (distal). f in collection.
Fibula (both epiphyses). J
The above list shows that the epiphyses of the limb bones tend to fuse first to
the bones nearest the body — humerus and femur — and the fusion proceeds outwards
to the digits. In the youngest animal (1892. 11. 7.1) the separate elements of the
vertebrae have not yet fused, and the fusion of the vertebral epiphyses to the centra
in the older animals appears to start at the cervical end of the column and proceed
caudally.
In order to see how the limb bones and pelvis increase in length with age,
measurements were taken of the lengths of the pelvis and of the shafts of five limb
bones, not including the epiphyses, of three male animals of increasing age, and the
new-born specimen, as there can be no difference due to sex at this age. There are,
unavoidably, many inaccuracies in the table (Table IV). The shaft length is
difficult to measure exactly in the two youngest animals because of the large amount
of cartilage present where the epiphyses are not fully ossified. The proportions,
using the measurements of the largest specimens as 100%, are taken on the assump-
tion that all the animals will grow to this size. This is obviously not so, and even
this animal is not fully mature, but it is the oldest that is known to be male. The
table shows that the pelvis has over half (62.2%) of its growth in length to complete
after the animal is born. The corresponding figures for the limb bones are :
humerus 45.4% radius 50%, femur 54.3%, and tibia 55-4%-
THE MONK SEALS 251
Table IV
Registered
Humerus.
Radius.
Femur.
Tibia.
Pelvis.
No.
mm. %
mm. %
mm. %
mm. %
mm. %
M . monachus :
1892. 11. 7. 1
59 54'6
63 50-0
• 43 45'7
91 44-6
92 37-8
1894.7.27.3 c? .
76 70-0
80 65-1
■ 58 6i-7
127 62'2
147 60-4
1863. 4. 1. 1 6" •
101 93*5
. 118 93-6
. 88 93-6
l86 91- I
221 90-9
1894.7.27. 1 6*.
108 100
126 100
94 100
204 IOO
. 243 IOO
RELATIONSHIPS OF THE GENUS MONACHUS FLEMING 1822
Family Phocidae Gray, 1825.
Subfamily Monachinae Trouessart, 1904.
The Phocidae are divided into the subfamilies Phocinae, including Phoca, Erig-
nathus and Halichoerus, with the incisive formula 3 ; the Cystophorinae including
2
Cystophora and Mirounga, with incisors 2 ; the Lobodontinae, including Lobodon,
1
Hydrurga, Leptonychotes, and Ommatophoca with incisors 2 ; and the Monachinae
2
with the single genus Monachus also with incisors 2.
2
The Cystophorinae are quite distinct, and are not considered here. Some of the
characters which distinguish the Phocinae from the Lobodontinae are : the
zygomatic process of the maxilla with the posterior border subvertical, not extending
far backwards beneath the malar ; nails of all the digits well developed ; and the
outer digits of the pes not much prolonged beyond the others. In the Lobodontinae
the zygomatic process of the maxilla is prolonged backwards beneath the malar ;
the nails of the hind limbs rudimentary ; and the outer digits lengthened.
A comparison of members of the genus Monachus with members of the two
preceding subfamilies shows that in the skull the incisive formula is the same as in
the Lobodontinae, the extension backwards of the zygomatic process of the maxilla
is not quite so great as in the latter subfamily, but is very much greater than in the
Phocinae. All members of the Phocinae have large claws on both fore and hind
flippers, while Monachus agrees with the Lobodontinae in having the hind claws
very much reduced. A further similarity is in the shape of the hind flippers which
have the two outer digits very much longer than the inner ones, the first digit of the
fore flippers is the longest and the rest get gradually shorter. In the Phocinae the
fore flipper is much more square in shape and the digits are more nearly equal.
The Phocinae also differ in the position of their nostrils, which are on the anterior
end of the snout and more or less vertical. The nostrils of both the Monachinae
and the Lobodontinae are nearly on the dorsal surface of the snout and are almost
horizontal. As already noted in the various sections on the bones of the skeleton
of Monachus, these are in general much more like those of the Southern than the
252 THE MONK SEALS
Northern Phocids. The above evidence shows clearly that the Monachinae are
more closely related to the Lobodontinae than to the Phocinae.
ACKNOWLEDGMENTS
I have to thank Dr. G. von Wahlert of the Ubersee-Museum, Bremen, and Dr.
K. Zimmermann, Zoologisches Museum der Universitat, Berlin, for help in locating
the type specimen of Monachus schauinslandi and for information and photographs
relating to it ; and also Dr. F. R. Parrington of the University Museum of Zoology,
Cambridge for access to an articulated skeleton of M. tropicalis. I am very much
indebted to Dr. H. W. Parker, Keeper of Zoology, for his advice and particularly
for his assistance with the section relating to the growth of the skull. I am also
very grateful to Miss Hilary King and to Commander J. M. Chaplin, R.N. (Ret.) for
much help in the translation of papers.
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■ 1887a. The West Indian Seal [Monachus tropicalis Gray). Bull. Amer. Mus. Nat. Hist.
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— — 18876. The West Indian Seal. Science, 9 : No. 206, 35.
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— — 191 8. The Laysan Seal. Nat. Hist. N.Y. 18 : 399-400.
Alessandrini, A. 1819a. Osservazione su gl'inviluppi del feto della Phoca bicolor. Opuscoli
scientifici. 3 : 298-316.
18196. Bemerkungen iiber die Fotushiillen der Phoca bicolor. Arch. Physiol. 5 : 604-616.
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THE MONK SEALS
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256 THE MONK SEALS
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PLATE 3
Monachus monachus. Photographs of the specimen from Oran in the Jardin des Plantes,
Paris. The animal was blind in its right eye. Phot. J. E. K.
PRESENTED
11 JAN 1956
Bull. B.M. [N.H.) Zool. 3, 5
PLATE 3
MONA CH US MONA CH US
PLATE 4
Monachus monachus. a. From Hermann, 1779. b. From Buffon, 1782. c. From
Pennant, 1793.
Bull. B.M. (N.H.) Zobl. 3, 5
PLATE
MONA CHUS MONA CHUS
PLATE 5
Monachus tropicalis. Lateral view of skeleton 1887 . 8 . 5 . 1
hull. B.M. {N.H.) Zool. 3, 5
PLATE 5
MONA CHUS TROPIC ALIS
PLATE 6
M. schauinslandi. Skull of type specimen No. 32795, Zoological Museum, Berlin. A.
Dorsal view. b. Ventral view. c. Lateral view. d. Dorsal view of lower jaw. e. Lateral
view of lower jaw.
hull B.M. (N.H.) Zoot. 3, 5
PLATE 6
E.
D.
M. SCHAUINSLANDI
PLATE 7
M. tropical is. a. Fore limb.
Hind limb.
Bull. B.M. (N.H.) Zoo!. 3, 5
PLATE 7
M. TROPICAL! S
PLATE 8
Innominate bones of a. L. weddelli, b. M. monachus, c. H. grypus.
Bull. B.M. (N.H.) Zool. 3, 5
PLATE 8
L. WKDDELLI.
B C
M. MONACHUS. H. GRYPUS.
4g*»i
PRESENTED
11 JAN 1956
PRINTED IN GREAT BRITAIN BY
ADLARD AND SON, LIMITED,
BARTHOLOMEW PRESS, DORKING.
mT
2 2 FEB 1956
THE CEPHALOPODA OF
MADEIRA
RECORDS AND DISTRIBUTION
W. J. REES and G. E. MAUL
NOTES ON THE EUROPEAN
SPECIES OF ELEDONE
W. J. REES
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 3 No. 6
LONDON: 1956
THE CEPHALOPODA OF MADEIRA
RECORDS AND DISTRIBUTION
W. J. REES and G. E. MAUL
NOTES ON THE EUROPEAN SPECIES
OF ELEDONE
BY
W. J. REES
Pp. 257 — 293; Pis. 9-10 : 4 Text-Figures
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 3 No. 6
LONDON: 1956
THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, is
issued in five series corresponding to the Departments
of the Museum, and an Historical Series.
Parts will appear at irregular intervals as they become
ready. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.
This paper is Vol. 3, No. 6 of the Zoological series.
PRINTED BY ORDER OF THE TRUSTEES OF
THE BRITISH MUSEUM
Issued February, 1956 Price Sixteen Shillings
THE CEPHALOPODA OF MADEIRA
RECORDS AND DISTRIBUTION
By W. J. REES and G. E. MAUL
CONTENTS
I. Introduction .....
II. A Summary of Previous Records
III. Additions to the Fauna .
IV. An Analysis of the Cephalopod Fauna
i, The coastal fauna
2. The benthic slope fauna
3. The bathypelagic slope fauna
4. The pelagic surface fauna .
5. The epipelagic fauna
6. The oceanic bathypelagic fauna .
7. The abyssal fauna
V. Cephalopoda as Food of Alepisauvus ferox
VI. References .....
Page
259
260
261
276
276
276
276
276
277
277
277
278
280
SYNOPSIS
The cephalopod fauna of Madeira is revised and no fewer than 22 additional species are
added to the list, chiefly on account of the examination of the stomach contents of A lepisaurus
ferox Lowe, Aphanopus carbo Lowe and the sperm whale. A total of 18 species (many of them
rare and seldom taken in the nets of deep sea expeditions) are recorded from Alepisauvus ferox.
The following little known species have been found in the stomachs of sperm whales at Madeira :
Tetronychoteuthis dussumieri (Orbigny), Lepidoteuthis grimaldi Joubin and Architeuthis sp.
The composite fauna is analysed and compared with that of the Mediterranean.
I. INTRODUCTION
This preliminary paper on the cephalopod fauna of Madeira has its origins in an
investigation of the stomach contents of the voracious fish, Alepisauvus ferox Lowe.
The work at Madeira was pursued for some years prior to the despatch of all the
cephalopoda taken (together with representative local collections) to the British
Museum (Natural History) for identification. The material yielded many species
scarce in collections and it was deemed an opportune time to produce a new list
of Madeiran cephalopods.
This list is rich in the names of seldom-recorded species, chiefly because the
fishes Alepisauvus fevox, Aphanopus cavbo Lowe and the sperm whale have been
examined for stomach contents. The occurrence of the various rare species are
noted in the main part of the text, but special attention is drawn to the records of
Lepidoteuthis gvimaldi Joubin, Tetvonychoteuthis dussumievi (Orbigny), Histioteuthis
bonelliana (Ferussac) and Avchiteuthis from the sperm whale.
zool. 3, 6.
19
26o THE CEPHALOPODA OF MADEIRA
In this preliminary paper we have not given full descriptive accounts of the species
found ; this is reserved for further accounts of interesting species either jointly or
independently.
II. A SUMMARY OF PREVIOUS RECORDS
The first record of a cephalopod to be recorded in scientific literature from the
vicinity of Madeira was of a Taonid which d'Orbigny referred to Loligopsis pavo
Lesueur in the great Histoire des cephalopodes acetabuliferes (1835-1848) . It was later
pointed out by Rochebrune (1884) and Hoyle (1884) that d'Orbigny had confused
two species under this name and the Madeiran specimen (taken in sight of Madeira
by the naturalist Dussumier) was redescribed by the former under the name Phas-
matopsis cymoctypus.
Robert MacAndrew, who dredged extensively on the coast of Spain and Portugal,
both in the Atlantic and in the Mediterranean, does not mention a single cephalopod
from his dredgings at Madeira (1849-50).
White and Johnson, in their Handbook of Madeira (i860) list the following six
species :
Loligo vulgaris Lamarck.
Sepia officinalis L.
Octopus vulgaris Lamarck.
Octopus cuvieri (=0. macropus Risso).
Argonaut a argo L.
Ocythoe tuber culata Rafinesque.
In their second edition (1885) the fourth and last species are not listed. Hoyle
(1885, p. 69) confirmed the occurrence of Argonauta argo at Madeira in his Challenger
Report.
More species were added to the list by Girard (1892) who reported on the collections
of the Abbe Ernest Schmitz of the Seminario do Funchal, adding or confirming the
existence of the following at Madeira :
Loligo forbesi Steenstrup.
Spirula peronii Lamarck 1 (=Spirula spirula L.).
Sepia filliouxii Lafont (— S. officinalis).
Todarodes sagittatus (Lamarck).
Ocythoe tuber culata Rafinesque.
Octopus tuberculatus Blainville. 2
Girard was aware of the differences between L. vulgaris and L. forbesi and as the
species submitted to him by Schmitz proved to be L. forbesi, the former species was
dropped from the list ; nor was he able to confirm the existence of 0. macropus in
the area.
1 Apparently empty shells.
* Presumed here to be a juvenile specimen of 0. vulgaris.
THE CEPHALOPODA OF MADEIRA 261
Watson (1897) added no new species to the list of Madeiran cephalopods but
confirms that 0. vulgaris is a common species and states that Sepia officinalis is rare.
From 1890 onwards the various deep sea expeditions have taken cephalopods in
the N. Atlantic, sometimes near, sometimes some distance off Madeira, and the list
compiled by Nobre (1937) includes 20 species of which several have been taken a
considerable distance away and could not be termed local fauna. Some of the
species which he records were taken as far west from Madeira as the African coast
is to the east of it. The following species were taken too far away to be retained as
local Madeiran records :
Sepietta oweniana (Orbigny).
Onychia caribaea Lesueur.
Brachioteuthis riisei (Steenstrup) .
Histioteuthis bonelliana (Ferussac).
Taonius pavo (Lesueur) .
Japetella diaphana Hoyle.
As Girard's Octopus tuberculatus probably falls into the synonymy of 0. vulgaris,
Nobre's list becomes amended to 13 cephalopods known from the immediate vicinity
of Madeira. To these may be added Joubin's record of Leachia cyclura (1920).
III. ADDITIONS TO THE FAUNA
The Madeiran archipelago has a much richer cephalopod fauna as our investiga-
tions have shown, and we do not think we have by any means exhausted the numbers
of species which should occur off Madeira. A great many other cephalopods known
from the central North Atlantic remain to be reported from this area. Below is
given a list of 22 additional species which we are now recording from Madeira.
Heteroteuthis dispar (Ruppell).
Loligo vulgaris Lamarck.
Abraliopsis morisii (Verany).
Enoploteuthis leptura (Leach).
Pyroteuthis margaritifera (Ruppell).
Cucioteuthis unguiculata (Molina).
Tetronychoteuthis dussumieri (Orbigny). 1
Lepidoteuthis grimaldi Joubin. 1
Onychoteuthis banksi (Leach).
Architeuthis sp.
Histioteuthis bonelliana (Ferussac).
Ommastrephes pteropus Steenstrup.
Thysanoteuthis rhombus Troschel.
Chiroteuthis veranyi (Ferussac).
Mastigoteuthis schmidti Degner.
1 These two species are here recorded from the stomachs of sperm whales taken at Madeira. It is
hoped to provide a full description in another paper,
262 THE CEPHALOPODA OF MADEIRA
Taonius pavo (Lesueur) .
Cranchia scabra Leach.
Japetella diaphana Hoyle.
Eledonella pygmaea Verrill.
Vitreledonella richardi Joubin.
Alloposus mollis Verrill.
Tremoctopus violaceus delle Chiaje.
Family Spirulidae
Spirula spirula L.
Spirula peronii, Girard, 1892, p. 219 ; Watson, 1897, p. 318, Nobre, 1937, P- 5-
This species is known only from the dead shells at Madeira ; these are cast up
on the beaches of Porto Santo occasionally and are also known from Canigal (Girard) .
Bruun (1943) reporting on the living Spirula captured by the " Dana " expeditions,
believed that this species inhabits the continental slope, living at depths not
exceeding 1750 metres.
Family Sepiidae
Sepia officinalis L.
Sepia officinalis, White & Johnson, i860 ; Johnson, 1885 ; Watson, 1897, P- 3 J 6 ; Nobre, 1937,
P. 5-
Sepia fillouxii, Girard, 1892, p. 219.
Material seen :
1 specimen, dorsal mantle length of 205 mm. (Maul collection No. 144).
According to Watson (1897), Sepia officinalis is rare at Madeira, but the species is
very common all the year round at Madeira and adult specimens measuring about
25 cm. (in dorsal mantle length) are often seen at the local fish market. The species
goes by the name of Chouco and is eaten by the local population.
Family Sepiolidae
Heteroteuthis dispar (Riippell)
Material seen : All except No. 3281 from the stomachs of Alepisaurus ferox.
No. 3281, 1 mutilated specimen, dorsal mantle length 20 mm. from the
stomach of Aphanopus carbo, io.i.51.
No. 3365, 1 specimen, dorsal mantle length 25 mm., 10. iv. 1952.
No. 3490, 4 specimens, dorsal mantle lengths of 25, 28, 29 and 20 mm. — no
date.
No. 3802, 1 specimen, dorsal mantle length of 22 mm. — no date.
No. 3848, 2 specimens, dorsal mantle lengths of 8 and 12 mm. — no date.
This sepiolid has not previously been taken at Madeira, and although reported
from numerous stations in the Mediterranean (Degner, 1925). it has been found on
comparatively few occasions in the Atlantic. Each major expedition has, however,
THE CEPHALOPODA OF MADEIRA 263
secured specimens ; those found nearest to Madeira being those captured by the
" Talisman " (29 2' N., 14 49' N.) between the Canaries and the African coast
(Fischer & Joubin, 1907), and those of the " Thor " from just outside the Straits of
Gibraltar (Degner, 1925). Heteroteuthis dispar is often regarded as a deep sea
cephalopod, but it is difficult to justify this claim, which seems to be based on the
fact that the species has a luminous secretion. To quote Harvey (1952, p. 283) :
"It is sometimes said that the luminous secretion of deep sea Heteroteuthis has
replaced the ink of surface dwellers. This black fluid forms a poisonous smoke
screen which confounds the enemy while the squid makes a getaway. Although it
is not true that the luminous secretion has completely replaced the ink, it may
serve a similar purpose — to draw the attention of the enemy to a mass of lighting
material, while the squid moves off in another direction." However, an examination
of Degner' s details of capture in the Mediterranean and Atlantic shows that captures
were more frequent with only 300 metres of wire out than in deeper hauls, and it can
be reasonably assumed that the net was fishing at only half or less than half this
depth. The fact that Heteroteuthis has retained its ink sac is also an indication
that it can and does live above the threshold of light. Whether it can also live
below the threshold of light is so far unknown. As an item in the food of Alepi-
saurus ferox, it probably occurs at the depths at which this fish is taken on tunny
hooks at Madeira. It is also a regular item in the diet of the long-finned albacore
(Germo alalunga) in the Bay of Biscay (Bouxin & Legendre, 1936, p. 36), and it has
been reported from the stomach of the dolphin (Delphinus delphis) by Joubin
(1900, p. 10).
Sepietta oweniana (Orbigny)
Sepiola scandica, Fischer & Joubin, 1906, p. 204.
Sepietta oweniana, Bouxin & Legendre, 1936, p. 32 ; Nobre, 1937, P- 5-
Material seen :
No. 2812, 2 specimens, dorsal mantle lengths of 12 and 17 mm., Funchal
Harbour, 12. hi. 1943.
The " Travailleur " took this species in position 32 40' N., 18 54' W., some
distance to the south-west of Madeira. Its occurrence at Madeira is confirmed from
two specimens we have seen.
Family Loliginidae
Loligo vulgaris Lamarck
Material seen :
No. 2966, 1 juvenile, no data.
No. 3164, 5 juveniles from Funchal Harbour, 4. v. 1950.
No. 3848, 1 juvenile, dorsal mantle length of 7 mm. from stomach of Alepi-
saurus ferox.
The specimens from Funchal Harbour are juveniles of only 30-35 mm. in dorsal
mantle length. Loligo vulgaris is recorded as far north as the coast of Denmark,
264 THE CEPHALOPODA OF MADEIRA
but it is doubtful whether it breeds anywhere north of the latitude of Ushant at
the mouth of the English Channel. As a Lusitanian species it is well known in the
Mediterranean and on the Mauritanian coast, while recently Adam (1952) has demon-
strated that it occurs off the Congo and as far south as Algoa Bay. It has been
reported from the Canary Islands (Odhner, 1931) and, although said to occur at the
Azores (Drouet, 1858), its presence there requires confirmation.
Loligo forbesi Steenstrup
Loligo forbesii, Girard, 1892, p. 219 ; Nobre, 1937, P- 6.
No. 4355, Funchal Market 1 $ of 270 mm. in dorsal mantle length.
Girard based his identification of this species on many examples sent to him by
the Abbe Ernest Schmitz of Funchal. This species has often been reported from
the neighbouring African coast. Locally, Loligo goes by the name of Lula da costa,
but it is not known whether the two species are confused under this name. It is
very common in winter, and as an article of food stands next to 0. pteropus in
importance. Market specimens range from about 200 mm. to 500 mm. in dorsal
mantle length.
Abralia veranyi (Riippell)
Abralia veranyi, Berry, 1926, p. 257.
Abralia veranyi, Nobre, 1937, P- 7-
Material seen :
No. 2664, 1 specimen of 47 mm. dorsal mantle length, Funchal Harbour,
at night, 16.X.1941.
No. 2964, no data — 5 specimens, 37-50 mm. in dorsal mantle length.
No. 3514, Funchal Museum (no date).
This luminous squid is a Mediterranean-Atlantic species ; most of the records
of its occurrence are from the Mediterranean (Messina, Nice, Genoa, Naples and
Toulon), while in the Atlantic (apart from Madeira) it has been found at compara-
tively few places, viz : Cockburn Town, San Salvador Island (Adam, 1941) and off
the Congo (Adam, 1952).
These specimens provide additional records for Madeira. No. 3514, a male, has
the left ventral arm hectocotylized just as figured by Adam (1952, p. 63, fig. 25E).
This specimen has a dorsal mantle length of 42 mm. One of the female specimens
(Lot No. 2964) has a dorsal mantle length of 50 mm. ; this is considerably bigger
than Adam's largest specimens, which were up to about 40 mm. in dorsal length
(Adam, 1952, p. 70).
Abraliopsis morisii (Verany)
Material seen :
No. 3848, 6 specimens of 8-25 mm. in dorsal mantle length from the stomach
of Alepisaurus ferox.
THE CEPHALOPODA OF MADEIRA 265
This is the first time this small luminous squid has been recorded from Madeira
and from the stomach of Alepisaurus ferox. Records of this species from the
Mediterranean and the Atlantic are fairly numerous (see Chun, 1910).
Enoploteuthis leptura (Leach)
Material seen :
No. 2664, Funchal Harbour at night, 16.X.1941, 1 specimen.
No. 2963, Madeira, 1 specimen, no data.
No. 3487, Funchal Harbour at night, i.vi.1946, 1 specimen.
These are the first Madeiran records of this rather scarce cephalopod. It is known
from the African coast and Pacific (Ferussac & d'Orbigny, 1 835-1 848) and a juvenile
from the Atlantic (position o° 29' N., 18 57' W.) was captured by the Deutsche
Sudpolar-Expedition and figured by Chun (1910, Taf. xl, figs. 5 and 6) in the
" Valdivia " report.
Pyroteuthis margaritifera (Ruppell)
Material seen :
No. 2961, 2 specimens, dorsal mantle length of 35 mm., from stomach of
Alepisaurus ferox, March and April, 1944.
This is another well known Mediterranean- Atlantic, luminous cephalopod.
Joubin (1924, p. 52) reported it from the Azores but it does not seem to have been
recorded from Madeira, neither has it been recorded as an item in the diet of Alepi-
saurus ferox. The " Thor " took specimens in the Eastern Mediterranean with
200-300 metres of wire out, so that the species can be assumed to exist up to 100
metres of the surface.
Family Octopodoteuthidae
Cucioteuthis unguiculata (Molina)
Cucioteuthis unguiculata, Joubin, 1900, p. 51 ; Nobre, 1937, P- 6.
The " Princess Alice " took this species in position 32 32' 10" N., 19 24' 40" W.,
to the west of Madeira. There is a large brachial crown, in the Municipal Museum,
Funchal, from a specimen which was found dead floating at the surface near the
shore to the west of Funchal.
Onychia caribaea Lesueur
Teleoteuthis jattai Joubin, 1900, p. 67.
Teleoteuthis caribaea, Nobre, 1937, P- 7-
Nobre includes this species in his list of Madeiran cephalopods, but the position
at which this species was taken by the " Princess Alice " (1897) Cruise, St. 812,
31 04' N., 27 11/ W.) is nearly as far to the south-west of Madeira as these islands
are from the African coast. The species must therefore be omitted from the list.
266 THE CEPHALOPODA OF MADEIRA
Onychoteuthis banski (Leach)
Material seen :
No. 2455, no data, 1 specimen of 100 mm. in dorsal mantle length.
No. 3507, no data, 1 specimen of 130 mm. in dorsal mantle length.
No. 2965, no data, 1 poorly preserved specimen of 35 mm. dorsal mantle
length.
No. 3571, 1 specimen of 48 mm. dorsal mantle length, from stomach of
Alepisaurus ferox, 4 . xii . 1952.
No. 3848, 45 specimens, dorsal mantle lengths of 10, 13, 21-24 mm., stomach
of Alepisaurus ferox.
No. 3509, 1 mutilated juvenile, dorsal mantle length 24 mm., from the
stomach of Alepisaurus ferox.
Onychoteuthis banksi is here recorded for the first time from Madeira. No. 2965
is in poor condition and only provisionally referred to this species.
This is a well-known oceanic surface species which, in the neighbourhood of
Madeira, was taken in position 30 54' N., 24 11' N., by the Prince of Monaco in his
1912 cruise (Joubin, 1924, p. 48). Other nearby records for the " Terra Nova "
and the " Ara " are given by Rees (1949, p. 43) who drew attention to the habit of
" flying " in this species. New records of specimens being found on the deck of
ships are given by Adam (1952, p. 77).
Family Architeuthidae
Architeuthis sp.
A sperm whale, harpooned off Sao Lourenco on 12th June, 1952, vomited a squid
which had evidently just been swallowed because the latter still exhibited some signs
of life. This squid was large, with a weight of 150 kilograms, and could only have
been a species of Architeuthis. The following measurements were taken :
Overall length (i.e. including tentacles) . . 10,600 mm.
Dorsal mantle length . . . . .1,860
Length of tentacles
Length of arm
Circumference of mantle
Circumference of tentacles
8,500
2,800
1,360
330
Architeuthis was taken by the French steamer " Alecton " between Madeira and
Teneriffe in November, i860, and the species was subsequently given the name
Loligo bouyeri by Crosse and Fischer (1862, p. 138). A number of species of Archi-
teuthis have been described since Verrill's classical monograph and it is by no means
certain how many kinds there are. Fragments of the above specimen are kept in
the Museu Municipal, Funchal, and it is hoped to present a more detailed description
in another paper.
THE CEPHALOPODA OF MADEIRA 267
Family Histioteuthidae
Histioteuthis bonelliana (Ferussac)
Histioteuthis ruppelli, Joubin, 1900, p. 98.
Histioteuthis bonelliana, Nobre, 1937, P- 6.
Material seen :
No. 3905, 1 specimen, dorsal mantle length of 53 mm., from stomach of
Aphanopus carbo, 30 . x . 1953.
Nobre records this species as Madeiran on the basis of an arm found in position
41 o' N., 12 15' W., by the " Princess Alice " in 1894 (Joubin) ; this position is off
the north coast of Portugal, so that this could not be justified as a Madeiran record
of this species. Histioteuthis does occur, however, around Madeira, as recorded
above, and we are indebted to Mr. H. M. Sieyes for a photograph of a large brachial
crown of this species found in the stomach of a sperm whale captured off Sao Laurenco
lighthouse, Madeira on 12th June, 1952. It is interesting to note that this sperm
whale feeds on this species at Madeira, as at the Azores and elsewhere. The speci-
men from the stomach of Aphanopus carbo was much smaller, with an overall length
length of about 170 mm. It is noteworthy that Verrill (1880, p. 234) recorded
H. bonelliana from Alepisaurus ferox (as H. collinsii).
Family Brachioteuthidae
Brachioteuthis riisei (Steenstrup)
Tracheloteuthis riisei, Joubin, 1924, p. 75.
Brachioteuthis {Tracheloteuthis) riisei, Nobre, 1937, P- 6.
This wide-ranging surface cephalopod is included in the Madeiran fauna by Nobre
on the strength of a record of capture of this species in position 31 45' N., 20 17' W.,
by the " Princess Alice " (Joubin). This is some distance south and west of Madeira,
but such a widely distributed species is almost certain to be found in this area.
Family Ommastrephidae
Todarodes sagittatus (Lamarck)
Todarodes sagittatus, Girard, 1892, p. 220.
Ommastrephes sagittatus, Nobre, 1937, P- 8.
Material seen :
No. 3355, 1 juvenile, dorsal mantle length of 83 mm. from the stomach of
Alepisaurus ferox, 8 . iv . 1952.
No. 3356, 1 juvenile, dorsal mantle length 86 mm. from the stomach of
Alepisaurus ferox, 8 . iv . 1952.
No. 3363, 1 juvenile, dorsal mantle length 85 mm. from the stomach of
Alepisaurus ferox.
No. 4347, Funchal Market, i specimen of 185 mm. in dorsal mantle length.
268 THE CEPHALOPODA OF MADEIRA
Madeiran records in the literature are few and are based on the following speci-
mens : i $ taken by Schmitz, i $ by Nunes. To these may be added the specimens
from the stomach of Alepisaurus ferox noted above.
The sucker rings of the tentacles are mostly absent : the very long-sucker-bearing
region of the tentacle, extending for the greater part of its length, the almost smooth,
large tentacular rings and the presence of four rows of suckers near the tip of the
tentacles indicate that the species is T. sagittatus. Locally Todarodes is called
Cartucho ; it is extremely common on the fishing grounds in March, April and May.
It is also taken during other months, but it is rare at these times.
As a common cephalopod on the fishing grounds of the eastern North Atlantic
and the Mediterranean, Todarodes is preyed on by a number of animals, including
the dolphin, Delphinus delphis (Joubin, 1895, p. 32), Germo alahinga (Bouxin and
Legendre, 1936) and Gadus calliarias {Michael Sars Stations 7, 86, 352 and 115).
Joubin (1895, 1900, 1920 and 1924) also reported various ommastrephid fragments,
some of which probably belong to this species, from Polyprion americanum and
Grampus griseus.
Ommastrephes pteropus Steenstrup
Material seen :
Nos. 65 and 66, 2 fragments, Funchal Harbour, 16.iv.1941.
No. 193, 1 juvenile, dorsal mantle length 58 mm., 10. v. 1940.
No. 3443, 1 specimen, dorsal mantle length 270 mm. — no data.
No. 3678, 1 juvenile, dorsal mantle length 50 mm. from stomach of Alepi-
saurus ferox, 25 . iii . 1953.
D. W. Tucker Collection, No. 605, 1 juvenile, dorsal mantle length 80 mm.
D. W. Tucker Collection, Nos. 639-644, 6 half-grown specimens, gaffed
at night on Madeiran fishing grounds, August, 1953.
All these specimens have been referred to 0. pteropus ; some of them might equally
well have been referred to 0. bartrami as we find that the diagnostic key given by
Pfeffer (1912, p. 465) is unreliable and our specimens clearly belong to one species.
Adam (1952, p. no) also noted the unreliable nature of the connective apparatus
on the tentacles as a means of separating bartrami from pteropus. The number of
suckers, proximal to the nearest connective button, will vary as new buttons develop
with the growth of the tentacle, as indicated in one of our larger specimens where
there is a rudimentary button ; this points to the great need for a revision of these
species with more material than we have at our disposal.
The relationships of these two species to Ommastrephes caroli is equally puzzling.
In the species caroli the lateral membrane is a broad triangular flap in adult speci-
mens (Rees, 1950, pi. 1) while in younger specimens it is much narrower (see Hertling,
1938). Our specimens from Madeira (those of 2-3 ft. in total length) possess narrow
lateral membranes on the third arms which might conceivably develop into the
characteristic paper-thin flaps of caroli. As in the latter their chromatophores are
closely set and a deep chestnut-brown in colour,
THE CEPHALOPODA OF MADEIRA 269
Ommastrephes pteropus is very abundant during the summer months of July,
August and September and is extensively used for baiting the tackle for Aphanopus
carbo, tunny and other fish. It is found in great numbers in locally caught Germo
obesus. Large quantities of this squid, which goes under the local name of
Pota de limao or Lula de limao, are sold in the fish market for consumption by
the local people, who greatly appreciate it as food. The large, lemon-coloured
dorsal mark has been observed to flare up with a bright light when the animal
is gaffed.
Ommastrephids :
No. 2957, 1 juvenile, dorsal mantle length of 50 mm. from stomach of Alepi-
saurus ferox, March-April, 1944.
No. 3848, 3 juveniles, dorsal mantle length of 17, 20 and 25 mm. from the
stomach of Alepisaur us ferox.
No. 193, 1 juvenile, dorsal mantle length of 65 mm. from the stomach of
Alepisaurus ferox, 10 . v . 1940.
These fragmentary specimens are in poor condition and we have not been able to
identify them.
Thysanoteuthis rhombus Troschel
Material seen :
1 specimen, dorsal mantle length of 350 mm., taken at Funchal.
This is the first record of this powerful surface species from Madeira. Some years
ago a school of about 20 came close inshore, and this is the only specimen it was
possible to save for examination. Little is known about its distribution, but it
occurs in the Mediterranean (Naef, 1 923-1 928) and has been reported from the Cape
of Good Hope by Barnard (1934). It is likely, however, to be a cosmopolitan warm-
water species, as Sasaki (1929) has recorded it from a number of localities in Japanese
waters.
Chiroteuthis veranyi (Ferussac)
Material seen :
No. 3685, 1 Doratopsis larva of 80 mm. in overall length. Coll. A. A. Nunes,
8.vii.i948.
This larva was taken at the Pontinha at the surface at night in the light of an
electric lamp suspended over the water.
It is a well-known Mediterranean- Atlantic species whose larva is frequently taken
at places like Messina and Villefranche, where there is upwelling from deep water.
The closely related species Doratopsis exophthalmica was taken by the " Valdivia"
in position 31 59' N., 15 5' W. to the south-east of Madeira.
Published records of C. veranyi from the North Atlantic are few (see Adam, 1952,
p. 111), but to judge from the number of specimens recovered from the stomachs
of germon fished in the Bay of Biscay the species is not as rare as the absence of
records suggests (Bouxin and Legendre, 1936).
270
THE CEPHALOPODA OF MADEIRA
Mastigoteuthis schmidti Degner
Material seen :
1 specimen, dorsal mantle length of 107 mm., from the stomach of Alepi-
saurus ferox, December, 1944.
There are no previous records of Mastigoteuthis from Madeira and this specimen
is only provisionally referred to M. schmidti because we recognize the need for a
complete revision of the large number of species described from the Atlantic. As
regards the proportions of mantle and fin the species is very similar to M. schmidti.
Family Cranchiidae
Leachia cyclura Lesueur
Leachia cyclura, Joubin, 1920, p. 68.
Material seen :
No. 3513, 1 specimen, overall length 145 mm. from stomach of Alepisaurus
ferox — no date.
No. 3848, 1 juvenile, dorsal mantle length of 23 mm. from stomach of Alepi-
saurus ferox.
Leachia cyclura was taken by the Prince of Monaco on the fishing grounds of
Madeira in 1901 (Stn. 1235, 8th September, 32 34' N., 17 45' W.). This species
has rarely been captured in the Atlantic, and since it was described from the Atlantic
coast of N. America by Lesueur (1821), it has been recorded by Joubin only from
Madeira, the Azores and the Sargasso Sea. It is known from other oceans also.
Cranchia scabra Leach
No. 34970, 1 specimen, overall length of no mm. from stomach of Alepisaurus
ferox.
No. 34976, 1 specimen, Funchal Harbour, at night.
No. 3497c, 1 specimen, overall length 105 mm. from stomach of Alepisaurus
ferox.
This species is well known from the sub-tropical and tropical areas of the Atlantic
and other oceans, but does not appear to have been reported from Madeira. It is
also recorded for the first time from the stomach of Alepisaurus ferox.
There is no particular reason for supposing that this species is a deep-water
animal. Its presence in the upper 200 metres has been proved by the Deutsche
Sudpolar Expedition, which took specimens at 20 and 40 metres depth. It should
be noted that records from 2,000 and 3,500 metres by the " Valdivia " were made
with open nets (Chun, 1910).
THE CEPHALOPODA OF MADEIRA 271
Taonius pavo (Lesueur)
Taonius pavo, Joubin, 1900, p. 106 ; Nobre, 1937, P- 8.
Material seen :
No. 3902, 1 specimen without tentacles, partly macerated, from the stomach
of Alepisaurus ferox, 26 . xi . 1953.
This specimen agrees favourably with the specimen figured by Joubin (1900, p. 106,
pi. viii and ix), but the colour is of a deep reddish purple. Our specimen has a
dorsal mantle length of 275 mm. and a total length of 385 mm.
This example, from the stomach of A. ferox, is the first to be found at Madeira.
The specimen taken by the Prince of Monaco in 1897 (Stn. 817, position 30 42' N.,
27 32' W. was found well out in the Atlantic and a considerable distance from
Madeira.
Taonius pavo has all the appearance of being a bathypelagic species, and we have
wondered whether the occurrence of this squid (and also some others, notably,
Mastigoteuthis schmidti and Eledonella pygmaea) in this fish can be explained by
local up welling of deep water bringing these species near the surface.
Taonius cymoctypus (Rochebrune)
Loligop sis pavo, Orbigny, 1835-1849, p. 321 (pars).
Phasmatopsis cymoctypus Rochebrune, 1884, p. 15, pi. 1.
Taonius pavo, Girard, 1892, p. 220 ; Nobre, 1937, P- 8.
The only specimen known of this cephalopod was taken within sight of Madeira
by the French naturalist Dussumier and was confused with T. pavo by d' Orbigny.
Later Rochebrune demonstrated that it was a distinct species. A new description
of this curious squid is greatly needed.
Order OCTOPODA
Family Bolitaenidae
Japetella diaphana Hoyle
Japetella diaphana, Thore, 1949, p. 23, fig. 14.
Bolitaenella diaphana, Nobre, 1937, P- 3-
Material seen :
No. 2958, 4 specimens, total length ca. 50 mm., stomach of Alepisaurus ferox,
March- April, 1944.
No. 3357, 1 specimen, total length ca. 50 mm., from stomach of Alepisaurus
ferox, 8.1V.1952.
No. 3495, 2 specimens, total length of 50 and 70 mm., stomach of Alepisaurus
ferox.
No. 3496, 1 specimen, total length 70 mm., ? stomach of Alepisaurus ferox.
272 THE CEPHALOPODA OF MADEIRA
No. 3569, 2 specimens, total lengths 55 and 80 mm., stomach of Alepisaurus
ferox, 4.XH.1952.
No. 3594, 1 specimen, total length 65-70 mm., stomach of Alepisaurus ferox,
i.iii.1953.
No. 3903, 1 damaged specimen, total length 85 mm., stomach of Alepisaurus
ferox, 26.x. 1953.
There are numerous records of this pelagic octopod in the North Atlantic, in the
triangle, Canaries, Gibraltar, Azores (Thore, 1949, fig. 14, p. 23), but the first positive
records from Madeiran fishing grounds are given above.
Bouxin & Legendre (1936, p. 12) reported finding this species (one specimen in
each of three stomachs) in the germon (Germo alalunga) captured in the Bay of
Biscay. It is now recorded for the first time from Alepisaurus ferox and appears to
be a regular item in the diet of this fish.
For the Bolitaenella diaphana of Nobre, see Eledonella pygmaea.
Eledonella pygmaea Verrill
Eledonella diaphana, Joubin, 1900, p. 37, pi. 2, figs. 5-7.
Material seen :
No. 3800, 1 $ total length 150 mm., from the stomach of Alepisaurus ferox, no
date.
Eledonella pygmaea is less common than the preceding species. It has not
previously been recorded from Madeira or from the stomach of Alepisaurus ferox.
The " Princess Alice " (Joubin, 1900) took this species between Madeira and the
African coast and there are more recent records of specimens taken by the " Dana "
to the west and north of the Madeiran archipelago (Thore, 1949, fig. 41, p. 49).
Family Vitreledonellidae
Vitreledonella richardi Joubin
Vitv eledonella alberti Joubin, 1924, p. 38 ; Nobre, 1937, P- 3-
This species had been included in a list of Madeiran cephalopods by Nobre (1937),
who quotes Joubin's record of a specimen taken to the N.W. of Madeira (position,
33 40' N. to 33 52' N., 19 W. to 19 16' W.) by the Prince of Monaco.
Family Octopodidae
Octopus vulgaris Lamarck
Octopus vulgaris, White and Johnson, i860 ; Johnson, 1885 ; Girard, 1892, p. 218 ; Watson,
1897, P- 2 9° «' Nobre, 1937, P- 3-
Octopus tuberculatus , Girard, 1892, p. 218 ; Nobre, 1937, P- 4-
Octopus rugosus, Robson, 1929.
THE CEPHALOPODA OF MADEIRA
273
Material seen :
British Museum :
B.M. No. 1898. 5. 10. 2, Porto Santo, Madeira, Coll. H. S. Wellcombe (Robson,
1929, as 0. rugosus).
B.M. No. 1912.12.31.118-119, Porto Santo, Madeira, 60 fms., R. Kirk-
patrick (Robson, 1929, as 0. rugosus).
Funchal Market from mouth of Conger conger, 18 . viii . 1953, Coll. D. W. Tucker.
Funchal from a tide pool, 28. viii. 1953, Coll. D. W. Tucker.
Funchal Museum :
No. 2460, 1 post-larva.
No. 140, 1 specimen, dorsal mantle length 90 mm.
No. 3287, from the stomach of Aphanopus carbo.*
The common octopus, which goes by the local name of polvo, is very common all
the year round at Madeira and is on sale in the fish market. The fact that it is eaten
by the local population may account for the scarcity of specimens available for
examination. It is found, too, at the Cape Verde Islands, the Canaries and the
Azores. Juvenile octopods mentioned under the name Octopus tuberculatus seem
to belong to 0. vulgaris.
Octopus macropus Risso
Octopus cuvieri, Girard, 1892, p. 219 ; White & Johnson, i860 ; Nobre, 1937, P- 4-
Material seen :
Funchal Museum, —
No. 39, 1 $, total length 475 mm., Camara de Lobos, 30. xi. 1940.
No. 139, 1 <J, total length 1,010 mm. — no date.
No. 2460, 2 juveniles in alder i stage — no date.
No. 3390, 1 J, total length 550 mm. — no date.
No. 3493, 1 juvenile in the alderi stage, from the stomach of Alepisaurus ferox.
This species was originally listed as occurring at Madeira in White & Johnson's
Handbook, but the name was omitted in the second edition (1885). Subsequent
authors have not recorded new material, so that evidence of its occurrence at
Madeira was needed. The new records given above dispel this doubt and confirm
its presence at Madeira.
Octopus sp.
Material seen :
No. 2960, 1 partly-digested specimen with a total length of 120 mm., from
the stomach of Alepisaurus ferox, March-April, 1944.
No. 3504, 1 specimen from stomach of Conger conger (L.), 18.vii.1952.
No. 3577, 1 specimen with a dorsal mantle length of 14 mm., from stomach of
Alepisaurus ferox, 10 . xii . 1952.
It has not been possible to positively identify these specimens with either of the
foregoing species or any other, because of their immaturity and poor condition.
* From the stomach of Aphanopus carbo according to a fish gutter
zool. 3, 6. 20
274 THE CEPHALOPODA OF MADEIRA
Family Alloposidae
Alloposus mollis Verrill
Material seen :
No. 2460, 1 ?, total length 115 mm., March- April, 1944.
No. 2960, 1$, total length 115 mm., stomach of Alepisaurus ferox.
No. 3491, 5 ?, total lengths of 80, 115, 115, 115 and 117 mm., from stomach
of Alepisaurus ferox, no date.
No. 3494, 1 ?, total length 175 mm., stomach of Alepisaurus ferox, no date.
No. 3804, 1 $, total length 115 mm., stomach of Alepisaurus ferox, 12 .vi. 1953.
No. 3848, 1 $, total length of 50 mm., stomach of Alepisaurus ferox , no date.
This species has not been previously recorded from Madeira, but has been taken
at the Azores and other places in the North Atlantic, mainly in proximity to coastal
waters. Thore, who plotted its distribution (1949, fig. 69, p. 72), regards it as a
cosmopolitan, tropical and sub-tropical coast-loving species.
The series now reported from Madeira are all small females (50-175 mm. in total
length, compared with VerruTs specimen of 812 mm.) (Verrill, 1882). The paucity
of records of Alloposus are noted by Thore, who suggests that either it is a rare
animal or is able to evade plankton nets or spends " relatively short periods of its
life-cycle in open waters, then soon returning to a life at the bottom, especially on
the continental slopes." These new records suggest that the species is by no means
as rare as the literature suggests and that we lack the gear to take it in numbers.
At the smaller sizes (50-175 mm.) it seems to be a regular item in the diet of Alepi-
saurus ferox.
Family Tremoctopodidae
Tremoctopus violaceus delle Chiaje
Material seen :
No. 3404, 1 9, total length 480 mm.
No. 3406, 1 9, total length 510 mm.
No. 3408, 1 9, total length 305 mm.
There are no previous records of Tremoctopus from Madeira. It is well known
from the Mediterranean but records from the North Atlantic are remarkably few
(these are summarized by Robson, 1932, and Salisbury, 1953).
Family Ocythoidae
Ocythoe tuberculata Rafinesque
Ocythoe tuberculata, Girard, 1892, p. 218 ; Robson, 1932, p. 202.
Oxythoe tuberculata Nobre, 1937, P- 4-
Material seen :
1 9, British Museum, No. 1858. 3. 31. 6 (Robson, 1932).
No. 2460, 1 9, total length 115 mm. (33 mm. DML), no data.
THE CEPHALOPODA OF MADEIRA
275
No. 3440, i 9, dorsal mantle length of 190 mm. — no data.
No. 3441, 1 ?, dorsal mantle length of 180 mm. — no data.
No. 3492, 1 ?, dorsal mantle length of 50 mm., from the stomach of Alepi-
saitrus ferox.
No. 3848, 1 ?, dorsal mantle length of 12 mm., from the stomach of Alepi-
saurus ferox.
Girard records having seen a specimen of this species in the Museu do Seminario
do Funchal and there is another in the British Museum.
There are two large specimens noted here (Nos. 3440 and 3441) ; They compare
favourably in size with one reported by Berry (1916) from California. Large as
these are, Robson (1932, p. 205) records having seen one of 280 mm. (DML) at
Monaco.
The young ? specimen (No. 3492) has a total length of 195 mm., and is remarkably
well preserved. The tubercules, connected by ridges, on the ventral side of the
mantle, are well developed and the aquiferous pores prominent. In another, even
smaller specimen (No. 2460), these characteristics are easily seen.
Ocythoe tuberculoid is another cosmopolitan species in tropical and sub-tropical
waters of all oceans ; as far as we know, it is essentially a species which frequents
the surface layers. Bouxin & Legendre (1936, p. p. 31) found it in the stomach of
the germon in the Bay of Biscay, while Joubin (1900, p. 26) reported three females
from the stomach of Grampus griseus taken off Monaco.
It is now recorded for the first time as an item in the diet of Alepisaurus ferox.
Family Argonautidae
Argonauta argo L.
Argonauta argo, Girard, 1892, p. 218 ; Hoyle, 1885, p. 69 ; Watson, 1897, p. 274 ; Nobre,
1937- P- 5-
Material seen :
No. 3489, 1 juvenile $, dorsal mantle length 37 mm., no data.
Early records from Madeira and Porto Santo are noted by Girard and Hoyle
respectively, and it is not surprising that this cosmopolitan, warm-water species
should be found here.
A juvenile female (without shell) is provisionally referred to this species. Watson
(1897, p. 274) records one perfect specimen from Porto Santo, and by this we assume
(as he was a conchologist) a shell or brood chamber.
Females of about 20-100 mm. in dorsal mantle length are found in great quantities
in the stomachs of Alepisaurus ferox at Madeira during the months of March, April
and May. There is one record for February and another of a single specimen in
November. The shells are usually crushed, but sometimes they are quite intact.
Eggs have been seen in shells of not more than 50 mm. in diameter. There is, in
addition, a single record of a small female (of about 35 mm. in shell diameter) caught
in Funchal Harbour at night. Large empty shells are not uncommonly found
washed up on the beach of Porto Santo.
276 THE CEPHALOPODA OF MADEIRA
IV. AN ANALYSIS OF THE CEPH ALOPOD-FAUN A
Surrounded as it is by deep water, the Madeiran Archipelago has a composite
fauna with elements from different habitats. In general, the fauna may be said
to be a Mediterranean- Atlantic one, which we group as follows :
1. The coastal fauna
This includes both littoral and continental shelf species, viz : Sepia officinalis,
Sepietta oweniana, Loligo vulgaris, Loligo forbesi , Todarodes sagittatus, Octopus vulgaris
and 0. macropus.
All these are common also in the Mediterranean and eastern Atlantic. Only the
two species of Octopus have a wider distribution in the Atlantic and elsewhere.
2. The benthic slope fauna
No Madeiran representatives of this fauna have yet been taken. The deep-water
octopods of the genus Bathypolypus are typical examples of species found on the
bottom below the 200 m. line.
3. The bathy pelagic slope fauna
In the grouping of these species we are on less sure ground because of the scarcity
of locality and depth records. Despite this we believe that the normal habitat of the
adults of the species enumerated below is close to the bottom on the continental
slope below the 200 m. line, with a lower limit perhaps at 2,000 m.
Spirula spirula.
Lepidoteuthis grimaldi.
Cucioteuthis unguiculata.
Tetronychoteuthis dussumieri .
Architeuthis sp.
Histioteuthis bonelliana.
Ommastrephes pteropus.
4. The pelagic surface fauna
By these we mean the species that are habitually found on or very close to the
surface. They include :
Ommastrephes pteropus (juveniles).
Onychoteuthis banksi.
Tremoctopus violaceus.
Ocythoe tuber culata.
Argonauta argo.
And possibly also
Thysanoteuthis rhombus.
THE CEPHALOPODA OF MADEIRA 277
5. The epipelagic fauna
In grouping pelagic forms we have to remember that many species have epipelagic
larvae while the adults may live well below the threshold of light. Up welling from
deep water on some coasts and near oceanic islands sometimes brings charac-
teristically deep-water species to the surface and it is not always possible to reach
positive conclusions.
The following oceanic species appear to live in the upper 500 metres :
Heteroteuthis dispar.
Abralia veranyi.
? Enoploteuthis leptura.
Pyroteuthis margaritifera.
Chiroteuthis veranyi (? juveniles only).
Leachia cyclura (juveniles).
Cranchia scabra.
Japetella diaphana (juveniles).
Alloposus mollis (some part of its life history).
6. The oceanic bathy pelagic fauna
These include deep-water species which live at or below the threshold of light (ca.
700 m.) and which are not confined to the immediate vicinity of the continental
slope. These species often have larvae which are epipelagic.
Chiroteuthis veranyi (adult).
Mastigoteuthis schmidti.
Leachia cyclura (adult).
Taonius pavo.
Japetella diaphana.
Eledonella pygmaea.
Vitreledonella richardi.
We think that the occurrence of these species in the Madeiran list is probably due
to the fact that there is upwelling which brings them into the upper waters or into
the hunting grounds of Alepisaurus.
7. The abyssal fauna
Truly abyssal forms like V ampyroteuthis infernalis Chun have not been taken at
Madeira.
In drawing up these lists we have paid but little attention to early records made
with open nets from deep water. All too often when a plankton haul has been made
with an open net from great depths (say 3,000 m.) to the surface, the resulting catch
is reported as from 3,000 metres.
It has been clear for some time, too, that among the Cephalopods, larvae and
adults do not always have the same vertical distribution, so that an animal may be
a temporary inhabitant of several distinct faunal zones during the course of its life
278
THE CEPHALOPODA OF MADEIRA
history. Concerning vertical migration or diurnal rhythms in Cephalopods, we
have no evidence, nor do we know how tolerant they are of changes in salinity. We
do know, however, that some species are sensitive to changes in temperature and that
a rapid lowering of temperature by two or three degrees is sufficient to kill some
forms (e.g. Architeuthis and Sepia officinalis).
The above is in broad outline a reasonable grouping of the species, but more
exact limits may be possible when we know much more about this group. It must
be admitted, for instance, that it is not easy to decide whether the distinction
between a bathypelagic slope fauna and an oceanic bathypelagic fauna can be
maintained.
As the Madeiran Archipelago is the nearest oceanic group of islands to the Mediter-
ranean, it may be pertinent to note that the following species in the list do not occur
there :
Slope animals.
Spirula spirula.
Lepidoteuthis grimaldi
Cucioteuthis unguiculata.
Enoploteuthis leptura.
Tetronychoteuthis dussumieri.
Architeuthis spp.
Bathypelagic species.
Mastigoteuthis schmidti.
? Leachia cyclura.
Taonius cymoctypus.
Taonius pavo.
Cranchia scabra.
Japetella diaphana. 1
Eledonella pygmaea.
Vitreledonella richardi.
Only two of the species, which may be termed slope and bathypelagic species,
Histioteuthis bonelliana and Chiroteuthis veranyi, are found, and are known to breed,
in the western Mediterranean. In this area the chief captures have been reported
from places like Villefranche-sur-mer and Messina, that is, where upwelling from
deep waters occurs.
It is inappropriate here to discuss the cephalopods of the Mediterranean fauna at
length, but it can be said that it is poor in bathypelagic species ; this is in agreement
with what is known concerning the paucity of deep water species of other groups of
animals (Ekman, 1953, p. 369).
V. CEPHALOPODA AS FOOD OF ALEPISAURUS FEROX
Alepisaurus ferox Lowe is well known as a voracious fish, and it has been known as
a predator on squid since Verrill (1880, p. 234) described a brachial crown of Histio-
teuthis from a fish captured off Nova Scotia in 1879.
Examination of this series of 260 stomachs from Madeiran fishing grounds
has yielded many comparatively rare species of squids which are not normally
captured by plankton nets. The total of 18 species demonstrates that cephalopods
can be regarded as regular items in the diet of this fish :
1 " Dana " St. 1132, 36 , 10' N., 2° 46' W. in the Western Mediterranean near Gibraltar did yield
one larval diaphana of 12 mm. (Thore, 1949, p. 28). We concur with Thore that this juvenile specimen
was probably swept in through the Straits of Gibraltar in the easterly surface current.
THE CEPHALOPODA OF MADEIRA
279
List of Species Eaten
Number of
Max.
stomachs
Total
number in
containing
Species.
number.
one stomach.
cephalopods.
D.M.L.
Heteroteuthis dispar .
8
4
4
8-29
Loligo vulgaris .
1
1
1
7
Abraliopsis morisii
6
6
1
8-25
Pyroteuthis margaritifera
2
2
1
35
Onychoteuthis banksi .
6
4
3
10-48
Todarodes sagittatus .
3
1
3
83-86
Ommastrephes pteropus
1
1
1
50
Ommastrephids .
5
3
3
17-65
Leachia cyclura
2
1
2
23-50
Cranchia scabra
2
1
2
39-40
Taonius pavo .
1
1
1
275
Japetella diaphana
13
4
8
50-85
Eledonella pygmaea .
1
1
1
90
Octopus macropus
1
1
1
22
Octopus sp.
1
1
1
15
Alloposus mollis
9
5
5
30-40
Ocythoe tuberculata
2
1
2
12-50
Argonauta argo (see text)
20-100
The dorsal mantle length has been taken as a standard measurement for size,
and as a rough guide this is about half the total length of the animal in compact
species like Heteroteuthis dispar, Abraliopsis morisii, Pyroteuthis margaritifera,
Cranchia scabra, Japetella diaphana, Eledonella pygmaea, Alloposus and Ocythoe.
It represents about only a third of that of Histioteuthis bonelliana, where the body
is small and the arms disproportionately large. In the long and typical squids
like Loligo, Onychoteuthis, Todarodes, Leachia and Taonius, mantle length is usually
more than half the total length.
Japetella diaphana, Alloposus mollis and Heteroteuthis dispar are the species most
frequently represented and are perhaps rather sluggish creatures. It is particularly
noticeable that among the swift-moving species the specimens of Onychoteuthis
banksi were rather small, several being still in the planktonic stage.
As will have been noted already, none of the cephalopods from the stomach of
this fish can be regarded as truly deep-water species in the sense that this term can
be applied to forms like V ampyroteuthis infernalis. All might reasonably be expected
to occur in the upper 200 metres — the depth at which the fish were caught.
Of the eighteen species, only four, namely Loligo vulgaris, Todarodes sagittatus,
Japetella diaphana, Octopus macropus and Argonauta argo are common species in
collections. Of the remainder there is little doubt that they are also common species,
although records of capture are few. It is only recently that Japetella diaphana has
been demonstrated to be one of the most abundant pelagic species of cephalopod
(Thore, 1949), and Joubin (1933, p. 41) has similarly indicated from a preliminary
examination of the " Dana " collections that the Cranchiidae (to which Leachia
2 8o THE CEPHALOPODA OF MADEIRA
cyclura, Cranchia scabra and Taonius pavo belong) are also very numerous. Records
of Ocythoe tuber culata (and possibly also Alloposus mollis to a lesser degree) are very
few and do not imply scarcity but rather that the fishing gear hitherto employed by
expeditions is unsuitable for capturing animals of this group. The same difficulty
applies to the capture of swift surface species like Onychoteuthis banksi (Rees, 1949).
VI. REFERENCES
Adam, W. 1941. Cephalopoda. Mem. Mus. roy. Hist. nat. (2) Fasc. 21 : 83-161, pi. i-iv and
20 text-figs.
— ■ — 1952. C6phalopodes. Res. Sci. Exped. Oceanog. Beige Eaux Coheres Africaines de
I'Atlantique Sud, 3 (3) : 1, 142, pis. i-iii and 57 text-figs.
Barnard, K. H. 1934. Cephalopoda from the Cape of Good Hope. /. Conch. London,
20 : 44-45.
Berry, S. S. 1916. The Octopod Ocythoe in California. Pomona J. Entom. Zool. 8 (1).
1926. A note on the occurrence and habits of a luminous squid (Abralia veranyi) at
Madeira. Biol. Bull. Woods Holl, 51 : 257-268, 2 text-figs.
Bouxin, J., and Legendre, R. 1936. La fauna pelagique de I'Atlantique recueillie dans des
estomacs de germons au large du golfe de Gascogne — Deuxieme Partie : Cephalopodes.
Ann. Inst. Oceanog., Paris, 16 (1) : 1-99, 21 text-figs.
Bruun, A. F. 1943. The biology of Spirula spirula (L.). " Dana " Rep. 24 : 1-46, 2 pis.
and 13 text-figs.
Chun, C. 1908. Ueber Cephalopoden der deutschen Tiefsee — Expedition. Zool. Anz. 33 :
86-89.
1910. Die Cephalopoden. I : Oegopsida. Wiss. Ergeb. Tiefsee-Exped." Valdivia 18 :
1-402.
Crosse, H., and Fischer, P. 1862. Nouveaux documents sur les Cephalopodes gigantesques.
/. de Conch. Paris (3), 2 : 124-140.
Degner, E. 1925. Cephalopoda. Danish Oceanog. Exped. 1908-10, 11, C.l : 1-94, 52 text-
figs, and 3 charts.
Drouet, H. 1858. Mollusques marins des lies Acores. Mem. Soc. Agric. Sci., Belle Lettres,
Aude (II), 2.
Ekman, S. 1953. Zoogeography of the Sea. London: Sidgwick & Jackson, xiv -f 417 pp.
Ferussac, A. de, and Orbigny, A. d'. 1835-1848. Histoire naturelle generate et particuliere
des Cephalopodes acetabuliferes vivants et fossiles. Paris. 2 vols., text and atlas.
Fischer, H., and Joubin, L. 1906. Note sur les Cephalopodes captures au cours des expedi-
tions du " Travailleur " et du " Talisman." Bull. Mus. Hist. Nat. Paris, 1906 : 202-205.
1907. Cephalopodes. Exped. sci. " Travailleur " et du " Talisman," 8 : 313-353.
Girard, A. A. 1892. Les Cephalopodes des iles Acores et de l'ile de Madere. /. Cien
Math. Fis. Na. Lisboa, 1892.
Harvey, E. N. 1952. Bioluminescence. New York : Academic Press, xvi and 649 pp.,
187 text-figs.
Hertling, H. 1938, Ueber eine auf Juist gestrandete Sthenoteuthis caroli, (Furtado). Wiss.
Meeresuntersuch. 1 : 93-111.
Hoyle, W. E. 1884. On Loligopsis and some other genera. Proc. roy. Phys. Soc. Edin.
8 : 313-333-
1885. Report on the Cephalopoda. Rep. Sci. Res. H.M.S. " Challenger," Zool. 16 (44 :
1-245, pis. i-xxxiii and 10 text-figs.).
Joubin, L. 1895. Contribution a l'etude des Cephalopodes de I'Atlantique Nord. Res.
Camp. Sci. Monaco, 9.
1900. Cephalopodes provenant des campagnes de la " Princesse Alice " (1891-1897).
Ibid. 17.
THE CEPHALOPODA OF MADEIRA 281
1920. Cephalopodes provenant des campagnes de la " Princesse Alice " (1898-1910).
Ibid. 54 : 1-95. 16 pis.
1924. Contribution a l'6tude des Cephalopodes de l'Atlantique Nord. Ibid. 67 : 1-113.
14 pi. and 6 tables.
1933. Notes preliminaries sur les Cephalopodes des Croisieres du Dana (1921-1922).
4 C Partie. Ann. Inst. Oceanog., Paris, 13 (1) : 1-49, 48 text-figs.
Lesueur, C. A. 182 1. Descriptions of several new species of Cuttlefish. /. Acad. Nat. Sci.
Philad. 2 (1) : 86-101, 4 pi.
Lonnberg, E. 1896. Notes on some rare Cephalopods. K. Vetensk-Akad. Forhandl. 1896,
No. 8 : 603-612, 4 text-figs.
Mac Andrew, R. 1852. Note of the mollusca observed during a short visit to the Canary and
Madeira Islands, etc., in the months of April and May, 1852. Ann. Mag. Nat. Hist. (2),
10 : 100-108.
Nobre, A. 1937. Moluscos testaceos marinhos do Arquipelago da Madeira. Mem. Mus.
Zool. Univ Coimbra, 1937 ( x ) No. 98 : 1-101.
Odhner, N. H. 193 i. Beitrage zur Malakozoologie der Kanarischen Inseln. Lamelli-
branchien, Cephalopoden, Gastropoden, Arch. Zool. Stockholm, 23A (14) : 1-116, 2 pis.
and 48 text-figs.
Pfeffer, G. 191 2. Die Cephalopoden der Plankton Expedition. Ergeb. Plankton-Exped.
Humboldt-Stiftung 2, F,a : 1-8 15, text and atlas.
Rees, W. J. 1949. Note on the hooked squid, Onychoteuthis banski. Proc. Malac. Soc.
London, 28 : 43-45.
1950. On a giant squid, Ommastrephes caroli, Furtado, stranded at Looe, Cornwall.
Bull. Brit. Mus. Zool. 1 (2) : 31-41, 2 pis., 6 text-figs, and 3 maps.
Robson, G. C. 1929-1932. A Monograph of the Recent Cephalopoda, Parts I and II. London :
British Museum (N.H.).
Salisbury, A. E. 1953. Mollusca of the University of Oxford Expedition to the Cayman
Islands in 1938. Proc. Malac. Soc. 30 (1 and 2) : 39-54, pis. 7 and 8.
Sasaki, M. 1929. A Monograph of the dibranchiate cephalopods of the Japanese and adjacent
waters. /. Fac. Agric. Hokkaido Imp. Univ., Sapporo, 20, Suppl. No. 1-357, 3° pl s - an( i
159 text-figs.
Sverdrup, H. U., Johnson, M., and Fleming, R. 1946. The Oceans, their Physics, Chemistry
and General Biology. New York.
Thore, S. 1949. Investigations of the " Dana " Octopoda : Part I : Bolitaenidae, Amphi-
tretidae, Vitreledonellidae and Alloposidae. " Dana " Rep. 33 : 1-85, 69 text-figs.
Verrill, A. E. 1879-80. The cephalopods of the north-eastern coast of America. Part I :
The gigantic squids (Architeuthis) and their allies, with observations on similar large
species from foreign localities. Trans. Conn. Acad. Sci. 5 : 177-257 ; pis. 13-25.
1880-81. The cephalopods of the north-eastern coast of America. Part III : the
smaller cephalopods including the " squids " and the octopi with other allied forms. Ibid.
5 : 259-446, pis 26-56 and 3 text-figs.
Watson, R. B. 1897. On the marine mollusca of Madeira ; with descriptions of thirty-five
new species, and an index-list of all the known sea-dwelling species of that island. /.
Linn. Soc. 26 : 233-329, pis. 19 and 20.
NOTES ON THE EUROPEAN SPECIES
OF ELEDONE
WITH ESPECIAL REFERENCE TO EGGS AND LARVAE
By W. J. REES, D.Sc.
(British Museum (Natural History))
CONTENTS
Page
I.
Introduction .......... 283
II.
Eledone cirrhosa (Lamarck)
(a) Spawning
(b) The Egg Masses
(c) Description of the larvae
285
285
285
286
in.
Eledone moschata (Lamarck)
(a) Egg Masses and early stages
287
287
IV.
Distribution of the European Species
289
v.
The Differences between E. cirrhosa and E.
moschata
290
VI.
References ......
•
291
SYNOPS]
[S
The newly hatched larvae of the Lesser Octopus, Eledone cirrhosa (Lamarck), are described
and figured for the first time from material collected by the Fishery Board for Scotland.
Records of eggs and spawning have been brought together and additional ones listed. The
eggs and larvae of this species are compared with those of the Mediterranean E. moschata and
the distribution of both species reviewed. E. moschata occurs throughout the Mediterranean
and its distribution outside is limited to neighbouring coasts, north and south of the Straits of
Gibraltar. The records show that E. cirrhosa occurs in the western Mediterranean and along
the western coasts of Europe to Iceland, the Faroes and northwards to Trondhjem on the
Norwegian coast. The characteristics of the two species are compared.
I. INTRODUCTION
In this paper some new observations are made on the eggs and larvae of the Lesser
Octopus (Eledone cirrhosa Lamarck) and our previous knowledge of its reproduction
and distribution is summarized. Early naturalists confused this species with Eledone
moschata (Lamarck) and it has been deemed desirable to bring together what is known
of the eggs and larvae, as well as the distribution of both species to enable a summary
of the differences between the species to be presented. 1
Although Eledone cirrhosa is much more widely distributed in North European
waters than the Common Octopus (Octopus vulgaris Lamarck), it is surprising that
so little is known about its habits and life history. Its morphology and anatomy was
the subject of a memoir by Isgrove (1909).
1 The larva of a South African Eledonid, Pareledone nigra, has recently been described by me (see Rees,
1954)-
284
NOTES ON THE EUROPEAN SPECIES OF ELEDONE
Text-fig. i. — The recorded distribution of Eledone cirrhosa in inshore waters. Trawling
records from the central North Sea are not included. The sources of all the records are
given in the bibliography.
NOTES ON THE EUROPEAN SPECIES OF ELEDONE 285
II. ELEDONE CIRRHOSA (LAMARCK)
(a) Spawning
The spawning of Eledone cirrhosa in captivity was first noted by Joubin (1888)
who observed it in an aquarium at Banyuls in the month of June. According to
Joubin there were about 30 groups of eggs (and traces of another 30) most of them
being eaten by the female. Each bunch contained 5-19 eggs, the greatest number
being laid first. The eggs were white in colour, each being 7-8 mm. in length.
Spawning of Eledone was also noted by Gravely (1908) in an aquarium tank at
Port Erin, Isle of Man, in July. He noted that eggs were about 7 mm. by 2-5 mm. in
diameter and that one to four bunches of eggs were laid almost every day for about
a month, after which spawning was less regular and was soon over.
Isgrove (1909) states that one Eledone lays about 800 eggs and that these are
spawned in groups of 25-30 eggs. Egg clusters 4-7 cm. long were collected in Aberdeen
market by Bowman ; the full sized ova were 8-9 mm. in length and about 4 ■ o mm.
in diameter at the broad end (Russell, 1922). It now seems that these eggs are a little
too large to be those of E. cirrhosa, but their identity cannot be known for certain
until we have some information on the eggs of Graneledone verrucosa (Verrill) ; this
species replaces E. cirrhosa to the north of the Faroes.
Spawning in aquaria has been noted at Plymouth in January (Marine Biological
Association, 1931) and in July (Isgrove, 1909) ; at Port Erin in July (as noted by
Gravely) and in September (Moore, 1937, p. 196). A female captured on the Dogger
Bank spawned in the Heligoland aquarium in January (Hertling, 1936, p. 294)
Stephen (1944, p. 252) mentions several clusters of ova from N.N.W. of Ronas Voe,
Shetland (position, 6o° 42' N., i° 46' W.) trawled on 4th April, 1927, and, as the
embryos were well developed it can be assumed that spawning occurred in February
or early March.
From this it appears that Eledone may spawn all the year round, and this is
borne out by records of larvae, noted by Stephen (1944, p. 251). He observed that
although larvae were present all the year in the plankton catches of the Fishery Board
for Scotland, they were more frequent during the period May to August ; this
suggests maximum spawning in April, May, June and July. As to whether this
period of more intensive spawning is linked with the known seasonal migration of
Eledone into inshore waters we have insufficient evidence. Even the kind of haunt
chosen for spawning in nature is not known for certain, but it appears that Eledone
does not brood over its eggs, nor does it seem to lay them in shells or pots, as does
Octopus vulgaris, for it has never been taken with its eggs.
(b) Egg masses
Apart from eggs seen in aquaria, Eledone spawn is rarely taken and only two
positive records are known to me, the batches trawled near Ronas Voe in the Shet-
lands (Stephen, 1944) and a very large cluster from the Eddystone Grounds off
Plymouth in the collections of the Plymouth Laboratory.
The large egg mass from the Eddystone Grounds contains a very large number of
undeveloped eggs. The stalks of the eggs are very short and the largest eggs are 67
286 NOTES ON THE EUROPEAN SPECIES OF ELEDONE
mm. in length by 2-4 mm. in width. Some eggs are extremely small, being only
2-85-3-6 mm. in length by 1-0-1-5 mm. in diameter (PL 9).
The small egg-cluster from Ronas Voe contains embryos in an advanced stage of
development. The eggs themselves are 6-65-6-79 mm. in length by 2-94-3-29 mm.
in width. There is still a large yolk mass, but the embryos are well formed. Chroma-
tophores are developing the arms, head and body and the Kollikersche buschel are
clearly seen on the head and mantle (PI. 10, figs. 1-4). At this size (3 mm. in. ventral
mantle length) the single row of suckers on the subequal arms are formed. .
A third cluster in the British Museum is without any particulars ; the eggs are
poorly preserved and resemble those from Ronas Voe.
Records of larvae of E. cirrhosa are few. Lo Bianco (1909) found young Eledone
in the plankton in the Bay of Naples ; those found in April had a length (?
total length) of 40 mm. and those found in October a length of 120 mm. He gives no
adequate description and there is no certainty as to which species he had. Scottish
records of larvae are given by Russell (1922) and Stephen (1944). Russell noted that
the arms are much shorter in proportion to the body (3 : 5) in the young, the back is
smooth and covered with large chromatophores and the body is generally surrounded
by a thick, soft, translucent cuticle.
In the Bay of Biscay area Bouxin and Legendre (1936, p. 24) found seven specimens
ranging from 21-33 mm. in length in the stomachs of germon in positions approxi-
mately 90-100 miles to the south-west of Glenans and at about 250 miles to the
south-west (that is, near Cape Finisterre).
A new description is therefore needed and is given below.
(c) Description of the larvae
Dr. A. C. Stephen has kindly allowed me to examine a series of 18 larvae, from the
catches of the Fishery Board for Scotland, which were reported by him in 1944.
It has already been noted that larvae still in the eggs may reach a ventral mantle
length of 3 mm. in large eggs. In small eggs this length would presumably be less,
so that planktonic larvae of approximately this size can be regarded as having been
taken within a few days of hatching.
The young larva (PL 10, figs. 5-6) has much the same shape as the larva of
Octopus vulgaris (the only species it could be confused with in British waters) with its
short arms and rather squat mantle. In details, however, there are big differences,
the most noticeable feature being the arrangement of the chromatophores on the
mantle. Here the entire mantle is uniformly covered with large reddish brown
chromatophores, and these are also prominent on the head and arms. On the latter
there is a single row and those of the central portion of the head are deep-seated.
Overlying these in the transparent outer integument are other fainter reddish brown
chromatophores, which increase in number and size with growth. The outer integu-
ment of the eye is silvery and sometimes has a greenish hue. 1
The arms are subequal and do not have the thin cirriform tips seen in very young
larvae of Octopus vulgaris. No. 12 (Table I) has 6-7 fully formed suckers on each arm
1 All references to colour mean colour of preserved specimens in alcohol.
NOTES ON THE EUROPEAN SPECIES OF ELEDONE
287
with the clear rudiments of 8-9 more at the tip of the arm. A slightly larger specimen
(No. 13) has 9 suckers on each arm and about 8 rudiments at the tip.
The largest larva available (No. 19) has about 28 suckers plus rudiments at the tip.
The first proximal sucker has a diameter of 0-56, the second, third and fourth have a
diameter of 07, 0-84 and 1-05 respectively. Suckers 5-7 are the largest and then they
diminish in size distally. The web is subequal reaching to the 7th-8th suckers, that is,
to about one-third of the length of the tentacles. This specimen taken on 27th
November, 1930, was recorded from square E 13 b by Stephen (1944).
This late larva has developed a mantle fin-ridge as in the adult. The ventral
mantle is smooth but the dorsal mantle and head are both covered with tubercles.
The larval chromatophores are still discernible because of their large size, but the
areas in between have become covered with a large number of small chromatophores.
As noted above, the arms of the larval Eledone are quite short at hatching from
the egg and they are usually not so long as the ventral mantle (Table I). Growth of
the arms of the planktonic larvae is more rapid than mantle length (Figs. 3 and 4)
and is also reflected in the relation of mantle length to total length (Fig. 2) .
Table I.
— Larvae Collected by the Fishery Board J
or Scotland
No
A
r
I.
2.
3-
4-
5-
6.
7-
8.
9-
10.
Total length .
6-5
yo
6'45
7.9
8-95
8-25
8-2
8-12
8-34
7.84
Dorsal mantle length
(tc
>
eye) .
3*9
4-2
3-85
4-2
4'9
4*7
4-83
4.4
5-05
4*55
Ventral mantle length
3-6 5
3*35
2-52
3-85
4-06
3'8 5
4-06
3*43
4*2
3-64
Head width .
2-8
3-o
2-8
3'5
3'55
3-5
3-57
3'5
3*43
3'5
Mantle width .
3-15
3-2
3*55
3-85
3-78
3-64
4'4
3-99
3-64
4-2
Diameter of eye
o-8
1-05
1-05
1-05
1 -i
1 -2
I -2
1 -26
1-05
i-4
Length of arms
2-45
2-8
2'5
3.08
3-5
3-15
3'i5
3-15
2-8
2-8
Diameter of suckers
0'2
o-3
0-22
0-32
o*3
No
0.38
o-34
o-3
0-38
0-32
A
r
^
II.
12.
13.
14.
15-
16.
17-
18.
19.
Total length .
8.4
7*35
9'3I
10*08
7'7
9'45
9'87
T-7-T-5
29-0
Dorsal mantle length
(tc
)
eye) .
4-97
4-62
5-n
5'39
4-76
5*25
5-95
7'7
13-0
Ventral mantle length
3-92
3'7i
4 - 4 I
4-48
4-06
4*34
5-04
6-58
I2-0
Head width
3'5
3'7i
3 '99
4'55
3-78
4'i3
4*2
6-3
9-0
Mantle width .
■ 3-8 5
4-06
4-62
4*9
4'2
4-76
4-69
7-14
io-o
Diameter of eye
1 -26
1 -26
i- 4
i'4
i'3
1-6
i-5
2-2
3-5
Length of arms
■ 3*22
2-66
3-7i
3*92
2-94
3-85
3'8
8-4
16-0
Diameter of suckers
• o-3
0-32
0-4
0-4
o*3
0-4
0-4
0-7
I -12-
I -2
III. ELEDONE MOSCHATA (LAMARCK)
(a) Egg masses and larvae
We know less about the spawning and larval stages of this species than we do of
Eledone cirrhosa. Korschelt (1893) described an egg-mass found on a Pinna shell at
288 NOTES ON THE EUROPEAN SPECIES OF ELEDONE
— ~T T
i ■ ■
i i i i
i • i i i i 1 r 1 1 1 i
i i i r— t —
1
-
o
-
1 1 ' T 1
LENGTH
-
LU
H
. Z
<
o
"
.
o
„
°o'
^
0*°
TOTAL LENGTH
-
o
i i
•
1 1 1 1
i i i i i i i i i i i i
1 1 1 1 1
i
10 15 20 25 30
Text-fig. 2. — The relation of mantle length to total length in larvae of
Eledone cirrhosa. Measurements are in mm.
15
10
-| r — t - - » 1 — « I — '■ " ■ | — — r 1 « 1 1 1 « 1 ~» 1— — i T
O
h <
op o
o ^o
°8> °°
VENTRAL MANTLE LENGTH
J •— i ■ i t i i \ i i i l i i i i * u
2 4 6 8 10 12
Text-fig. 3. — The relation of arm length to ventral mantle length in larvae of
Eledone cirrhosa. Measurements are in mm.
NOTES ON THE EUROPEAN SPECIES OF ELEDONE 289
Rovigno in the Adriatic. He gave the total number of eggs as 65-70 ; this mass
was made up of small clusters of 2-4 eggs joined to the main mass by a common
stalk. The eggs (excluding stalk) were 15 mm. in length, that is, the same length as
the eggs figured by Jatta (1896, tav 7, fig. 3).
Korschelt was under the impression that he had the eggs of E. aldrovandi (i.e.,
cirrhosa) , but this mistaken view was corrected with the identification of the large
eggs with E. moschata by Jatta and confirmation by Gravely (1908) that the smaller
eggs belonged to E. cirrhosa. There have been embryological studies on development
°"T T" 1 J 1 — T 1
1
1-2
•
1-0
.
0-8
DIAMETER
O
0-6
0-4
SUCKER
<P*>
a
0-2
O O
VENTRAL MANTLE LENGTH
■
R
,
2 4 6 8 10 12
Text-fig. 4. — The relation of sucker diameter to ventral mantle length in larvae
of Eledone cirrhosa. Measurements are in mm.
in the egg (Sacarrao, 1943, 1945, 1951 and 1952) but there is no detailed description
of the newly hatched larva. Sketches of juvenile Eledone have been published by
Jatta (1896, tav 7, figs. 5 and 10), while Naef (1923) gives a drawing of a post-larval
E. moschata from Trieste. A new description is much needed.
IV. DISTRIBUTION OF THE EUROPEAN SPECIES
Eledone cirrhosa (Lamarck) is found in the Mediterranean, but its full distribution
there is not known. It appears to be a common species in the western part, but there
are no records for the eastern Mediterranean. Under the name Eledone aldrovandi
there are numerous records for the western Mediterranean (Jatta, 1896), but there
are no records east of Dalmatia (Robson, 1932, p. 266) and according to Ninni
(1884) it does not penetrate to the head of the Adriatic.
In the Atlantic E. cirrhosa has a much less restricted distribution than E. moschata.
zool. 3, 6. 21
ago NOTES ON THE EUROPEAN SPECIES OF ELEDONE
It is a common species on the European continental shelf extending to southern
Iceland, the Faroes (Brunn, 1945, p. 8) and the west coast of Norway. In the open
areas of the shelf it appears not to extend beyond the line Iceland-Faroes-Bergen, but
is a common species southwards of this line. On the Norwegian coast itself it reaches
Ostnesfjord in the Lofotens, but is said to be scarce north of the Trondhjemsfjord
(Grieg, 1933).
Its distribution in inshore British waters as recorded in the literature on cephalo-
pods is plotted on Map I, and, quite apart from the fact that such maps tend to
reflect areas worked by biologists, the species appears to be scarcer in the southern
North Sea than elsewhere. Records from trawling grounds offshore have not as a rule
been plotted (those from the North Sea trawling grounds are given by Grimpe, 1925),
and it has not been possible to search for all the occasional records buried in the
transactions of local natural history societies.
Eledone moschata (Lamarck) is a Mediterranean species which ranges from Istanbul
(Digby, 1949), the Syrian coast (Gruvel, 1931) and Palestine (Bodenheimer, 1937) to
the adjoining region of the Atlantic. Korschelt (1893, p. 68) implies that it is the
common Eledone of the Adriatic and it has also been found commonly at Naples
(Jatta, 1896, and Naef, 1923), the Gulf of Marseilles (Vayssiere, 1917) and Monaco
(Boone, 1933) to quote only a few of the more recent records. Outside the Mediter-
ranean the species appears to be rare and records are few. The " Talisman " took a
female specimen in the Bay of Cadiz from a depth of 60 m. (Fischer & Joubin
1907, p. 328), while Adam (1941, p. 140) reported the first specimen from the African
Coast (Baie du Levrier, Port-E'tienne, Mauritania).
There are no authentic records from northern Europe ; Nielsen's Eledone moschata
from the Faroes (1930) proved on re-examination to be E. cirrhosa (Brunn, 1945, p. 9).
V. THE DIFFERENCES BETWEEN ELEDONE CIRRHOSA
AND E. MOSCHATA
The main differences between the species are tabulated below.
Eledone cirrhosa Eledone moschata
No musk odour. . Musk odour.
Skin with warts on dorsum. . Skin smooth.
Ridge along edge of mantle. . No pallial ridge.
The seven non-hectocotylized arms of the . The seven non-hectocotylized arms carry a
male carry close-pressed, flattened suckers, double series of transverse lamellae at their
forming cirri at the tips of the arms. tips.
The two retractor muscles of the gills are . The two retractor muscles of the gills are
fused at their base and form a " Y " inserted separately.
Spermatophore with spines. 1 . Spermatophore without spines.
Colour of adult : Light yellowish brown with . Colour of adult : Greyish brown colour with
diffuse rust-brown patches on the back. darker almost black patches on the dorsal
Ventral mantle pale ivory or pinkish side. Preserved animal is grey to dark
yellow with a greenish iridescent sheen. grey with numerous dark patches.
Eggs, Pyriform, ca. 7 x 2-5 mm. in clusters . Eggs sausage-shaped ca. 15 x 4 mm. in
of about 30. clusters of two to four.
1 Fort (1941) creates a new genus Acantheledone for E. cirrhosa laying particular stress on this character.
NOTES ON THE EUROPEAN SPECIES OF ELEDONE 291
Eledone cirrhosa (cont.) Eledone moschata (cont.)
Newly hatched young ca. 3-0 mm. in ventral . Newly hatched young probably ca. 8 mm.
mantle length. in ventral mantle length.
Parasites : . Parasites :
Dicyemennea eledones (Wagner, 1857). D. eledones.
Chromidina coronata (Foettinger, 1881). D. moschatum Whitman, 1882.
VI. REFERENCES
Adam, W. 1933. Notes sur les cephalopodes. Ill : Les Cephalopodes du Sud de la mer du
Nord. Bull. Mus. Roy. Hist. Nat. Belg. 9, No. 46 : 1-45 Charts A-D.
1939. Notes sur les cephalopodes. X : Sur quelques cephalopodes de la mer d'Islande.
Ibid. 15, No. 16 : 1-13, 4 text-figs.
1 94 1. Resultats scientifiques des croisieres du Navire — licole Beige Merator.
Cephalopoda. Mem. Mus. roy. Hist. Nat. Belg. (2), Fasc. 21 : 83-162, pi. i-iv and 20 text-figs.
Bodenheimer, F. S. 1937- Prodromus faunae Palestinae. Mem. Inst. Egypte, 33 : 1-286,
4 text-figs.
Boone, L. 1933. Scientific results of cruises of the yachts " Eagle " and " Ara " 1921-1928,
William K. Vanderbilt, Commanding. Coelenterata, Echinodermata and Mollusca. Bull.
Vanderbilt Mus. 4 : 1-2 17.
Bouxin, J., and Legendre, R. 1936. La faune pelagique de l'Atlantique recueillie dans les
estomacs de germons au large du golfe de Gascogne. Deuxieme Partie : Cephalopodes.
Ann. Inst. Oceanogr. Paris 16 (1) : 1-99, 21 text-figs.
Bruun, A. F. 1945. Cephalopoda. Zoology of Iceland, 64 (64) : 1-15, 4 text-figs.
Chumley, J. 1918. The Fauna of the Clyde Sea Area. Glasgow.
Clarke, W. J. 1936. Yorkshire Cephalopoda. /. Conch. Lond. 20 (9) : 257.
Dautzenberg, P., and Fischer, P. H. 1925. Les mollusques marins du Finistere et en
particulier de la region de Roscoff. Trav. Stat. Biol. Roscoff, Fasc. 3 : 1-180.
Degner, E. 1925. Cephalopoda. Danish Oceanogr. Exped., 1908-10, II, C. 1 : 1-94, 52
text-figs.
Digby, B. 1949. Cephalopods from local waters at the University of Istanbul. Nature, 163,
No. 4141, p. 411.
Evans, W. 1899. Notes on the effects of the recent October gale upon marine life on the
coasts of the Lothians. Ann. Scott. Nat. Hist. 29 : 6-1 1.
Fischer, P. 1867. Catalogue des Nudibranches et Cephalopodes des cotes oceaniques de la
France. /. de. Conch., Paris, 15 : 1-15.
and Joubin, L. 1907. Cephalopodes. Exped. Sci. " Travailleur " et du " Talisman."
8 : 313-353-
Forbes, E., and Hanley, S. 1853. A History of British Mollusca. London : J. van Voorst.
Vol. IV.
Forrest, J. E., and Waterston, A. R. 1934. Scottish records of Cephalopods. I : A
" Cephalopod Year " in the Clyde. Scot. Nat. 1934 : 29.
Fort, G. 1937. Note sur les especes francaises du genre Eledone. Bull. Soc. Hist. Nat.
Toulouse, 71 (3) : 309-314.
1 94 1. Eledone Leach et Acantheledone n. g. deux genres fondes sur la structure des sperma-
tophores (Moll. C6phalopodes) C.R. Acad. Sci. Paris, 212 (17) : 724-726.
Godfrey, R. 1900. Eledone cirrosa in West Lothian. Ann. Scot. Nat. Hist. 1900 : 185.
Gravely, F. H. 1908. Notes on the spawning of Eledone and on the occurrence of Eledone
with suckers in double rows. Mem. Manchester Lit. Phil.-Soc. 53 (4) : 1-14, 2 pis.
Grieg, J. A. 1933. Cephalopods from the West Coast of Norway. Bergens. Mus. Aarb. 1933,
No. 4 : 1-25, pi. i-iv and 1 text-fig.
Grimpe, G. 1925. Zur Kenntnis der Cephalopoden fauna der Nordsee. Wiss. Meeresunter-
suchungen. Biol Anst. Helgoland, 16 (3) : 1-124, 1 pi. and 34 text-figs.
292 NOTES ON THE EUROPEAN SPECIES OF ELEDONE
Gruvel, A. 193 i. Les Etats de Syrie. Richesses marines et fiuviales. Exploitation actuelle,
453 PP-> 30 Pis.
Gyngell, W. 1928. Cephalopoda landed at Scarborough in 1927. /. Conch. Lond., 18 : 216.
1929. Cephalopoda etc. at Scarborough in 1928. /. Conch., 18 (10) : 287.
Haddon, A. C. 1886. Recent Contributions to the Marine Invertebrate Fauna of Ireland.
Zoologist (3) 10 : 1-7.
Haines, F. H.,1945. Notes on general zoology, 1944. Mollusca. Pap. Hampshire Field Club,
16 (2) : 233.
Hargreaves, J. A. 1910. Marine Mollusca of the Yorkshire Coast. /. Conch. Lond. 13 (4) :
99-105.
Hertling, H. 1936. Mitteilungen iiber Todaropsis eblanae (Ball), Octopus vulgaris L. und
Eledone cirrosa (Lam.) aus. der Nordsee. Zool. Anz. 114 (11-12) : 289-296, 1 text-fig.
Horsman, E. Additions to the marine fauna of the Aberystwyth district. Aberystwyth Studies,
4 : 259-269.
Hoyle, W. E. Report on the Cephalopoda. Rep. Sci. Res. H.M.S. " Challenger," Zool. 16
(44) : 1-245 pis. 1-33, 10 text-figs.
Isgrove, A. 1909. Liverpool Mar. Biol Comm. Memoir XVIII : Eledone.
Jatta, G. 1896. I cephalopodi viventi nel Golfo di Napoli. Fauna und Flora des Golfes von
Neapel. Monogr. 23 : 1-268, pi. 1-3 1 and 64 text-figures.
Jeffreys, J. G. 1869. British Conchology , London : J. van Voorst. Vol. V.
Joubin, L. 1888. Sur la ponte de YEledone et de la seiche. Arch. zool. exper. et gen. (2)
6 : 155-163, 1 text-fig.
1920. Cephalopodes provenant des Campagnes de la " Princess Alice " (1898-19 10)
3 e S6rie). Res. Camp. Sci. Monaco, 54 : 1-95, 16 pi.
Korschelt, E. 1893. Uber den Laich und die Embroynen von Eledone. Sitz. Ber. Get.
Naturf. Berlin, 1893, No. 2 : 68-73, 2 text-figures.
Lo Bianco, S. 1909. Notize biologiche riguardante specialmente il periods maturita sessuale
degli animali del Golfo di Napoli. Mitth. Zool. stat. Neapel, 19 : 513-761.
Marine Biological Association, 1931. Plymouth Marine Fauna. Second Edition.
Massy, A. L. 1909. The Cephalopoda Dibranchiata of the Coasts of Ireland. Fisheries Ireland
Sci. Invest. 1907, I (1909) : 1-39, PI. I— III.
1 91 3- Further records of the Cephalopoda Dibranchiata of the Coast of Ireland.
Ibid. 1912, V (1913).
1928. The Cephalopoda of the Irish Coast. Proc. Roy. Irish Acad. 33 {Section B),
(2) : 25-37.
Moore, H. B. 1937. Marine Fauna of the Isle of Man. Proc. and Trans. Liverpool Biol.
Soc. 50 : 1-293.
Naef, A. 1921-28. Fauna e Flora del Golfo di Napoli. Monogr. 35 : Die Cephalopoden.
Nichols, A. R. 1900. A list of the Marine Mollusca of Ireland. Proc. Roy. Irish Acad. 5
(4) : 477-662.
Nielsen, E. 1930. Cephalopoda. Zoology of the Faroes. Part 56.
Ninni, A. 1884. Catalogo dei Cefalopodi Dibranchis osservati nell'Adriatico. Att. Acc.Ven.-
Tren.-Instr. Padova, 9 : 159.
Nordga ard , . 1 92 3 . The Cephalopoda Dibranchiata observed outside and in the Trondh j em-
fjord. Kgl. Norske. Vidensk. Selsk. Shrift. 1922, No. 5 : 1-14. 4 text-figs.
Rees, W. J. 1954. The larva of Pareledone nigra (Hoyle) from South Africa. Proc. malac.
Soc. 31 : 50-51, 1 pi.
Rendall, R. 1931. Notes on Eledone cirrosa Lam. /. Conch. Lond. 19 : 161-162.
Robson, G. C. 1932. A Monograph of the Recent Cephalopoda. Part II : The Octopoda.
British Museum (Nat. Hist.), London.
Russell, E. S. 1922. Report on the Cephalopoda collected by the research steamer " Gold-
seeker " during the years 1903-1908. Fisheries Scotland Sci. Invest. 1921, III : 1-45, pis.
i-v.
NOTES ON THE EUROPEAN SPECIES OF ELEDONE 293
Sacarrao, G. F. 1943. Observations sur les derniers phases de la vie embryonnaire de
" YEledone." Arq. Mus. Bocage Lisboa, 14 : 25-35, 1 pi. and 3 text-figs.
1945- Etudes embryologiques sur les Cephalopodes. Ibid. 16 : 33-68, 9 text-figs.
1951. Notice on the embryonic shell sac of Octopus and Eledone. Ibid. 22 : 103-105,
PI. I.
1952. Quelques remarques sur une comparison des ontogeneses d'Octopus et d'Eledone.
Rev. Fac. Cienc. Lisboa (2) C, 2 (2) : 215-222.
Service, R. 1892. Eledone cirrosa Lamarck in the Solway Firth. Ann. Scott. Nat. Hist.
1892 : 202.
Stephen, A. C. 1934. Scottish records of Cephalopods. Scot. Nat. 1934 : 29.
— ■ — ■ 1944. The Cephalopoda of Scottish and adjacent waters. Trans. Roy. Soc. Edin. 61
(1), No. 9 : 247-270, 14 text-figs.
Stevenson, J. A. 1935. The Cephalopods of the Yorkshire Coast. /. Conch., London,
20 (4) : 102-116.
Thorson, G. 1946. Reproduction and larval development of Danish marine bottom inverte-
brates, with special reference to the planktonic larvae in the Sound (0resund). Medd.
Komm. Dansk. Fisk. — og Havundersog. Ser. Plankton, 4 (1) : 1-523, 199 text-figs.
Tomlin, J. R. le B. 1892. Notes on the Marine Mollusca of the North Wales coast with
complete lists of the recorded nudibranchs and cephalopods. /. Conch. Lond. 7 (1) : 25-31.
Tregelles, G. F. 1896. The Marine Mollusca of Cornwall. Ibid. 8 (6) : 191-200.
Vayssiere, A. Note sur la presence d'un bras supplementaire chez un Eledone moschata
Leach. /. de Conch. Paris, 63 : 123-125, 1 pi.
PlESENTtQ^
" a 1956
EXPLANATION OF PLATES
PLATE 9
Egg mass of Eledone cirrhosa (Lamarck) from the Eddystone Grounds, Plymouth.
■nil. B.M. {N.H.) Zoology 3, 6
PLATE 9
Egg mass of Eledone cirrhosa (Lamarck)
PLATE 10
Eggs and larvae of Eledone civvhosa (Lamarck) del. G. L. Wilkins.
Fig. i. — Portion of an egg cluster from Ronas Voe, Shetlands.
Fig. 2. — A well developed larva in situ.
Fig. 3-4. — Views of larvae removed from their egg capsules.
Fig. 5-6. — Side and ventral views of a planktonic larva from the catches of the Fishery
Board for Scotland.
hill. B.M. (.V.//.) Zoology 3, 6
PLATE 10
/
i-L
\J
I f
«i: " -; •
EM- • .'-S
fey ' ■•;■'
Eggs and larvae oi Eledone cirrhosa (Lamarck)
ENT -Q^
?&£ <f *
PRINTED IN GREAT BRITAIN BY
ADLARD AND SON, LIMITED
BARTHOLOMEW PRESS, DORKING
- 2 MAR 1956
THE MONOTYPIC GENERA
OF CICHLID FISHES IN
LAKE VICTORIA
P. H. GREENWOOD
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 3 No. 7
LONDON: 1956
THE MONOTYPIC GENERA OF CICHLID
FISHES IN LAKE VICTORIA
BY
P. H. GREENWOOD
(East African Fisheries Research Organization, Jinja, Uganda)
Pp. 295-333 ; 10 Text-figures.
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 3 No. 7
LONDON: 1956
THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, is
issued in five series corresponding to the Departments
of the Museum, and an Historical Series.
Parts will appear at irregular intervals as they become
ready. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.
This paper is Vol. 3, No. 7 of the Zoological series.
PRINTED BY ORDER OF THE TRUSTEES OF
THE BRITISH MUSEUM
Issued February, 1956 Price Ten Shillings
THE MONOTYPIC GENERA OF CICHLID
FISHES IN LAKE VICTORIA
By P. H. GREENWOOD, B.Sc.
CONTENTS
Introduction
Synopsis of Genera
Victoria
Genus Macropleuro
Generic charac
Diagnosis
Macropleuro dus bic
Synonymy
Description
Syncranium ai
Coloration anc
Ecology
Affinities
Study materia
Genus Platytaeniod
Generic charac
Diagnosis
Platytaeniodus dege
Synonymy
Description
Syncranium
Ecology
Affinities
Study materia
Genus Hoplotilapia
Generic charac
Diagnosis
Hoplotilapia retrode,
Description
Syncranium
Coloration
Ecology
Affinities
Study materia
Genus Paralahidoch
Diagnosis
Paralabidochromis i
Description
Affinities
Discussion
Summary .
Acknowledgments
References
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nd synonymy
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ociated musculature
:hromatism
distribution records
nd synonymy
alenger
distribution records
nd synonymy
[gendorf
distribution records
nov. .
%e sp. nov.
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<TG IN
Lake
Page
298
299
299
299
301
301
301
304
305
308
310
3ii
3ii
312
312
314
315
315
315
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317
3i8
3i8
319
319
319
321
322
322
3^3
324
324
326
327
327
328
328
329
329
332
333
333
ZOOL. 3,
298 MONOTYPIC GENERA OF CICHLID FISHES IN LAKE VICTORIA
INTRODUCTION
In his revision of the Lake Victoria Cichlidae, Regan (1922) recognized four endemic
monotypic genera, the species being Astatoreochromis alluaudi Pellegrin, 1904,
Hoplotilapia retrodens Hilgendorf, 1888, Platytaeniodus degeni Boulenger, 1906, and
Macropleurodus bicolor (Boulenger), 1906. Subsequent collections made by the
Cambridge Expedition (1930-1931) extended the range of A. alluaudi to include
the Lake Edward system and Lakes Kachira and Nakavali (Trewavas, 1933), but
provided no further distributional data for the other three genera. Since Astatoreo-
chromis alluaudi occurs beyond the Victoria system, it is preferable to delay revision
of this species until numerous specimens from the different localities can be examined.
Some notes on the osteology and possible phyletic relationships of Astatoreochromis
have been published already (Greenwood, 1954).
A fifth genus which, but for its geographical separation, would not have been
distinguished from Labidochromis of Lake Nyasa, is recorded for the first time.
The present study is based on specimens collected by and for the East African
Fisheries Research Organization during a field study of the Lake Victoria Cichlidae,
and on material in the collections of the British Museum (Nat. Hist.), Museum
National d'Histoire naturelle, Paris, and the Museo Civico di Storia Naturale,
Genoa. It forms the first part of a revision of the Lake Victoria Haplochromis
species flock.
Notes on counts and measurements
The counts and measurements used are as defined by Trewavas (1935), except that
" length of head " is measured directly from the posterior margin of the operculum
to the premaxillary symphysis. This method has been found to yield more consistent
results than measurements taken between verticals through the posterior tip of the
operculum and the level of the tip of the snout, along a line parallel to the longitudinal
axis of the body. Likewise, length of snout is measured directly.
Other measurements and counts are :
Depth of preorbital : measured from about the middle of the orbital rim of the
preorbital bone along a line which continues the radius of the eye at this point ; the
line approximately bisects the bone.
Interorbital width : the least width of the roofing part of the frontal bones,
between the eyes.
Lower jaw : measured directly from the angle to the symphysis.
Lateral line scale series : After last upper lateral line scale, proceed to the scale
of the lower lateral line next behind the transverse row that includes the last scale
of the upper lateral line and slopes downwards and forwards from it.
In addition to those defined by Trewavas. Depth of cheek : the greatest depth
measured vertically from the lower orbital margin to the lower edge of the adductor
mandibulae muscles. In most specimens this is virtually a measurement of the depth
of the scaled portion of the cheek.
MONOTYPIC GENERA OF CICHLID FISHES IN LAKE VICTORIA 299
Post-ocular part of the head : measured directly from the posterior orbital margin
to the posterior tip of the operculum.
SYNOPSIS OF GENERA OF THE H APLOCH ROMI S GROUP
OCCURRING IN LAKE VICTORIA
1. Anal spines three ............ 2.
Anal spines four or more ........ Astatoreochromis
2. Teeth of upper jaw in two or more series anteriorly, but only in a single (rarely
double) series laterally ........... 3.
Teeth of upper jaw in 2-5 series both anteriorly and laterally . . . .4.
3. Anterior outer teeth disproportionately longer than the adjacent lateral teeth,
slender, unicuspid and procurrent. . . Paralabidochromis gen. nov., p. 327
Anterior outer teeth forming a graded series with the lateral teeth, not forwardly
directed . . . . . . . . . . . Haplochromis
4. Outer teeth enlarged and stout, with obliquely truncated and inwardly directed
crowns ......... Macropleurodus, p. 299
Outer teeth of both jaws small, bi- or unicuspid ; inner teeth in broad bands anteri-
orly and laterally ........... 5.
5. Lower jaw broad and flat, tooth bands in both jaws of uniform breadth antero-
posteriorly ; those of lower jaw continued posteriorly on to the ascending part
of the dentary ......... Hoplotilapia, p. 319
Lower jaw stout, rounded anteriorly ; teeth aggregated anteriorly into two pyriform
bands, contiguous at the symphysis ..... Platytaeniodus, p. 312
Genus MACROPLEURODUS Regan, 1922
Bayonia Boulenger, 191 1 {nee. Bocage 1865), type species Bayonia xenodonta Blgr.
Macropleurodus Regan, 1922, Proc. zool. Soc, Lond. 189 ; type species : Haplochromis bicolor
Boulenger, 1906, Ann. Mag. nat. Hist. (7) 17, 444.
Haplochromis (part) ""]
Paratilapia (part) > For references see synonymy under species.
Hemitilapia (part) J
Generic characters and synonymy
Prior to Regan's revision of the Lake Victoria Cichlidae (Regan, 1922) specimens
of the genus here recognized as Macropleurodus had been described as belonging to
several genera and species. When Regan [op. cit.) defined the apparently new genus,
Macropleurodus he did not have at his disposal specimens covering the wide range
necessary to determine ontogenetic changes in certain characters. Consequently he
failed to recognize that his genus was identical with Boulenger's Bayonia
(Boulenger, 1911), and incorrectly assigned the type specimens of this and one other
synonymous "species" to Haplochromis.
Regan's diagnosis of Macropleurodus gave particular emphasis to the posterior
premaxillary teeth :
. . . several inner series of small teeth anteriorly and three or four series of
enlarged teeth laterally, which are exposed when the mouth is shut" (Regan, 1922).
ZOOL. 3, 7. 22§
300 MONOTYPIC GENERA OF CICHLID FISHES IN LAKE VICTORIA
The two specimens on which this description was based have the posterolateral
inner teeth not only enlarged, but also similar in form to adjacent outer teeth. It
is now known, however, that in small individuals these inner teeth are only slightly
enlarged, if at all, and do not closely resemble the outer teeth. Specimens available
indicate that most fish between 80 and 90 mm. standard length have a dentition
intermediate between juvenile and adult types. In these, many of the typically
juvenile, stoutly bi- and unicuspid inner teeth are replaced laterally by teeth
differing only slightly in size and form from those of the outer series. Individuals
below 80 mm. lack enlarged inner teeth.
Thus, although Regan's diagnosis is incisive for fishes over 90 mm. S.L., it is not
sufficiently comprehensive to include smaller individuals.
On the other hand, the present collection shows that the morphology of the outer
teeth is equally diagnostic and, moreover, is little affected by the size of the individual.
Basically, the outer series is composed of stout teeth, having enlarged and obliquely
truncated anterior cusps and disproportionately smaller posterior cusps. In the
upper jaw these teeth are implanted obliquely to the long axis of the premaxilla,
so that their crowns lie at an angle to it and the anterior cusp points inwards. Teeth
of this type are present in all the specimens examined and are not known to occur
in any Haplochromis or related species.
The peculiar form of the outer teeth in Macropleurodus at once suggests affinity
with Bayonia xenodonta Boulenger (191 1), in which the outer teeth are described as
having " ... very large compressed crowns, with long anterior cusp directed
inwards and very short or indistinct posterior cusp ... ".In this respect, the
published figure of B. xenodonta (Boulenger, 191 1 and 1914) is somewhat misleading
since the teeth are stouter than depicted.
Through the courtesy of Dr. D. Guiglia, re-examination of the type specimen of
B. xenodonta has been possible and has confirmed that the outer teeth are alike in
Bayonia and Macropleurodus. Further, as Regan and Trewavas (1928) first observed,
Boulenger's description of the inner teeth is inaccurate. Posteriorly these teeth
form a double series, with six teeth on either side nearly as large as the outer, and
in this respect conform to the dental pattern of young Macropleurodus. Also, the
more anterior teeth of the two inner series are not " minute and conical " but are
in fact tricuspid. Thickening of the buccal mucosa (probably a fixation artefact)
has buried the inner teeth so that only their major cusps protrude. Two types of
tricuspid teeth are present : one, the usual small tricuspid tooth found in many
species of Haplochromis ; the other, stout and with the minor cusps displaced so as
to form a triangular crown having the major cusp at its apex. Similar trigonid
teeth are also found in Macropleurodus, both young and adult, but have not been
observed in Haplochromis.
There is also agreement in morphometric characters and, although Bayonia
xenodonta has only twenty-four outer teeth in the upper jaw, this number is within
the known lower range for Macropleurodus (see below).
There seems little doubt, therefore, that in all salient features the type and unique
specimen of Bayonia xenodonta is identical with small specimens of Macropleurodus
bicolor, and that the two are conspecific.
MONOTYPIC GENERA OF CICHLID FISHES IN LAKE VICTORIA 301
One other name has been given to cichlid fishes from Lake Victoria with teeth of
the Macropleurodus type, namely Hemitilapia mater famili as Pellegrin, 191 3. I have
examined the holotype (no. 12.278 in the collection of the Museum d'Histoire
naturelle, Paris) and found it to agree closely with Macropleurodus of similar size.
The teeth and dental pattern of the type and three other specimens represent an
advanced stage in the transition from juvenile to adult condition ; that is, the
postero-lateral premaxillary teeth of the first inner series are nearly comparable in
size and form with the adjacent outer teeth. The remaining inner teeth are small
and unequally bicuspid.
Regan (1922) tentatively synonymized H. materfamilias with Haplochromis
obliquidens Hilgendorf ; apparently he did not examine the type of H. materfamilias
{op. cit., pp. 157 and 158) and was misled by Pellegrin's description. Comparison
of the holotype with specimens of H. obliquidens at once reveals the existence of
fundamental differences in the dentition of the two species. In H. obliquidens the
movably implanted outer teeth are fine and numerous (50-70 in upper jaw) ; the
posterior cusp is wanting, except very occasionally in small fish, whilst the anterior
cusp, although obliquely truncate, is compressed and slender. In contradistinction,
the immovable outer teeth of H. materfamilias are coarse and less numerous (24-40) ;
with few exceptions a posterior cusp is present and the stout anterior cusp is
circular in cross section.
Diagnosis
Cichlid fishes of the Haplochromis group as defined by Regan (1920, 1922) but
differing from Haplochromis in having stout outer teeth with inwardly directed and
obliquely truncated crowns ; anterior cusp long, slightly decurved and not com-
pressed, the posterior cusp small and indistinct. Fishes over 90 mm. S.L. have one
or more inner premaxillary tooth-series composed laterally of enlarged teeth similar
in form to the adjacent outer teeth. Consequent upon the enlargement of the lateral
teeth, the dentigerous surface of the premaxilla is broader laterally than anteriorly.
In small individuals, where the inner teeth are small and bi- or unicuspid throughout
the series, the outer teeth are already characteristic. Teeth usually exposed laterally,
even when the mouth is shut.
Macropleurodus bicolor (Boulenger) 1906
(Fig. 1)
Haplochromis bicolor (part) Blgr. 1906, Ann. Mag. nat. Hist. (7) 17, 444 (type only).
Paratilapia bicolor (part) Blgr. 1907, Fish. Nile 479, pi. lxxxix, fig. 1 ; Idem, 191 1, Ann. Mus.
Genova (3) 5, 68 ; Idem, 1915, Cat. Afr. Fish. 3, 346, fig. 234.
Bayonia xenodonta Blgr. 191 1, Ann. Mus. Genova (3) 5, 70 ; Idem, 1915, op. cit., 488, fig. 338.
Hemitilapia materfamilias Pellegrin, 1913, Bull. Soc. zool. France, 37, 313 ; Boulenger, 1915,
op. cit., 492.
Haplochromis obliquidens (part), Regan 1922, Proc. zool. Soc, London, 188.
Macropleurodus bicolor, Regan, 1922, op. cit., 189.
302 MONOTYPIC GENERA OF CICHLID FISHES IN LAKE VICTORIA
Fig. i.—Macropleurodus bicolor, holotype. 7/8 N.S. (From Boulenger, Fishes of the Nile.
• •
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Standard Length (mm.)
MONOTYPIC GENERA OF CICHLID FISHES IN LAKE VICTORIA 303
35-
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Standard Length (mm.)
Fig. 2. — Scatter-diagram to show individual variation and allometry of the characters
indicated. The isometric relationship of head length and standard length is usual, as is
the allometry of head length and the inter-related snout, eye and preorbital measurements .
The negative allometry between length of lower jaw and headlength isunusual and seems
to be related to the stronger jaws and enlarged teeth of larger specimens. These diagrams
show scatter but not frequency. Within the size-range 100 to 125 mm. S.L., each dot in
the denser aggregates represents at least two specimens.
3 o6 MONOTYPIC GENERA OF CICHLID FISHES IN LAKE VICTORIA
5, a). Posteriorly, the neurocranium does not differ significantly from the generalized
type, except that the pharyngeal apophysis is not greatly depressed below the
parasphenoid, and its pro-otic buttress is broad and bullate.
Icm
Fig. 4. — Macropleurodus bicolor. (a) Neurocranium in left lateral view ; (b) dentary,
(c) premaxilla, both in right lateral view. Skeleton prepared from a specimen of
115 mm. S.L.
MONOTYPIC GENERA OF CICHLID FISHES IN LAKE VICTORIA 307
B
Fig. 5. — Haplochromis michaeli. (a) Neurocranium, (b) dentary, left lateral view ; (c)
dentary, occlusal view ; to show the skull and jaws of a generalized Haplochromis from
Lake Victoria. Skeleton prepared from a specimen of 115 mm. S.L.
The premaxilla shows considerable departure from the basic type found in most
Haplochromis species. Whereas in Haplochromis there is slight ventral arching of
the horizontal limb of this bone, in Macropleurodus the arch is greatly exaggerated,
with its point of maximum curvature at the broadest part of the tooth band (Fig.
4, b). Individuals with a marked dextrally developed tooth pattern have a cor-
responding degree of asymmetry in the two halves of the premaxilla, which is then
more acutely arched on the right side.
ZOOL. 3, 7. 22§§
308 MONOTYPIC GENERA OF CICHLID FISHES IN LAKE VICTORIA
A comparative study of jaw musculature was made with Haplochromis sauvagei
and H. prodromus, species in which the jaws show some deviation from the generalized
Haplochromis condition towards that of Macropleurodus. Negligible differences
were observed in the musculature of the three species, except that in M. bicolor the
adductor mandibulae I measures only 28% of the head length, whilst in H. sauvagei
and H. prodromus this muscle is 33% of head length. Adductor mandibulae II is also
relatively shorter in M. bicolor (21-0% compared with 33-6% in H. sauvagei and
26-9% in H. prodromus). In all other respects the cranial musculature of Macro-
pleurodus is typical for a generalized Haplochromis species.
Functional significance of the skull form, together with that of the well developed
teeth, jaws and muscles, is best considered in relation to the predominantly molluscan
diet of M. bicolor, and the manner in which its prey is secured. Whereas in most
Haplochromis species the lower jaw is somewhat obliquely inclined when the mouth
is shut, in M. bicolor it is almost horizontal. The position of the lower jaw at rest
directly affects the manner in which the whole mouth is protruded ; in typical
Haplochromis, maximal protrusion is achieved as a result of the mandible moving
from an inclined to a horizontal position. On the other hand, when the mouth of
M. bicolor is opened, mandibular movement is from the horizontal to a point below
it. Premaxillary movement is also directed downwards by the near vertical ethmoid
complex over which the premaxilla slides. In consequence, the whole mouth is
protruded ventrally, with the upper jaw slightly in advance of the lower. The short
and broad adductor mandibulae muscles allow the mouth to be retracted with
remarkable rapidity, thereafter mounting a powerful and sustained pressure on any
object held between the teeth. The crushing power of the jaws is further enhanced
by the stout outer teeth and well developed bands of lateral and postero-lateral inner
teeth.
Aquarium observations show ^hat M. bicolor, when feeding, usually approaches a
snail from above, rapidly protruding the mouth in an attempt to snatch its prey
from the substrate. Once the snail is firmly held and suitably orientated — generally
with the foot directly orally — there follows a series of short biting movements which
crush the shell and thus free the soft parts, which alone are ingested.
Attention has already been drawn to the similar skull structure in M. bicolor and
H. prodromus ; it is not surprising therefore to find that both species have similar
feeding habits.
Ontogeny. Alizarin preparations of larval M. bicolor, H. macrops (a generalized
species), and H. prodromus have been compared. These specimens reveal no funda-
mental differences in osteology or dentition of the three species when compared at
morphologically equivalent developmental stages. For example, at the latest stage
examined (9-0 mm. total length ; yolk sac almost completely resorbed) the small
conical outer teeth are morphologically and numerically identical in all three species.
From these admittedly few observations it would seem that characteristic adult
skull form and outer teeth must develop during post-larval ontogeny.
Coloration and polychromatism
Coloration in life. Adult females with greenish-yellow ground colour, becoming
MONOTYPIC GENERA OF CICHLID FISHES IN LAKE VICTORIA 309
lighter or silver ventrally. Dorsal and anal fins yellow-green, the former with or
without two irregular, dark, longitudinal stripes, the latter with two to four ill-
defined yellow spots on the posterior part. Caudal and ventral fins generally colour-
less, though the latter may sometimes be slightly dusky. Young of both sexes and
sexually inactive males have similar coloration.
Adult males (breeding coloration). Dark slate-blue ground colour, lighter, some-
times silver, ventrally. Chest, branchiostegal membrane, lower jaw and ventral
aspects of the cheek, black. Operculum and flank sometimes with a faint scarlet
flush. Dorsal fin dusky, with deep red spots and streaks between the rays, especially
intense on the soft part. Anal dusky, with well marked scarlet ocelli. Ventral fins
black. Intensity of male coloration is correlated with sexual state ; inter-grades
are known between the coloration described above and that of typical female or
juvenile coloration.
Besides normal sexually dimorphic coloration, certain fish exhibit a third colour
pattern, in the form of an individually variable piebald, black on a yellow-green
ground. The holotype is such a specimen (Fig. i). The bicolor pattern is clearly
composed of vertically arranged irregular and often interrupted dark bands, which
are generally continued across the body on to the vertical and paired fins. Although
some are more intensely blotched than others, no intergrades are known between
normal female coloration and bicolor variants.
With two exceptions all bicolor individuals examined were females. The colour-
pattern and degree of pigmentation differ in the two exceptional male fishes. Since
protandry might be suspected, the gonads were sectioned and examined micro-
scopically. In both fishes, however, there was evidence only of testicular tissue. In
one fish the pattern is typical ; in the other the pattern is less intense and occurs
on a darker ground than is typical for female fishes.
Accurate frequency-estimates for bicolor individuals are difficult to obtain, since
collectors show marked sampling bias in favour of these strikingly coloured fishes.
However, in more rigorously controlled collections from one area, bicolor frequency
amongst female fish in the 105 mm. to 125 mm. size class is approximately 30%,
an incidence sufficiently high to justify regarding the phenomenon as being due to
polymorphism and not to the maintenance of an atypical phenotype by recurrent
mutation.
If the two female colour patterns are accepted as an example of polymorphism, it
is necessary, ex hypothesi, to consider the selective balance which must exist between
the two forms. This question is further complicated by the apparently almost
completely sex-limited polymorphism in M. bicolor.
Since the genetical basis of polymorphism and sex determination is unknown
for Macropleurodus, some hypothesis at least is desirable before considering the
question of selective values for the two colour patterns.
If, as in many fishes, the female is the heterogametic sex, then a possible (and
doubtless oversimplified) explanation for this sex-limited polymorphism is that the
gene or gene complex underlying development of a bicolor pattern may lie in a sex
chromosome, be recessive to the gene or genes for normal colour, and be linked with
a recessive lethal gene. Thus full expression of bicolor pattern could only be manifest
3 io MONOTYPIC GENERA OF CICHLID FISHES IN LAKE VICTORIA
in the heterogametic sex. Males carrying the double complement of recessive bicolor
genes necessary for phenotypic expression in that sex, would, on the linkage supposi-
tion, die as a result of simultaneously receiving the two recessive lethal genes. Since
linkage is sometimes broken, a small percentage of male bicolor individuals might
well be expected and the two bicolor male fishes in this collection are possibly such
individuals.
By this reasoning, either selection in favour of polymorph genes must be sufficient
to compensate for loss of males and consequent unbalance of the sex-ratio or,
alternatively, the unbalanced sex ratio may be the factor preventing spread of bicolor
genes throughout the species, should these have a selective value slightly higher
than " normal ".
Two possible advantages associated with bicolor patterns, or genotypes, present
themselves. Firstly, a bicolor pattern is, in effect, a disruptive one and may thus
provide some protection against the attacks of predators. Studies on fish-eating
birds (Cott, 1952) and on piscivorous fishes such as Bagrus and Clarias (personal
observation) neither support nor negate this possibility, since M. bicolor has not
been found among the prey of these animals. Secondly, there is the possibility, also
unproven, that a female bicolor genotype, or a male heterozygous for bicolor, may
possess some physiological advantage over other genotypes.
Regrettably, then, insufficient positive evidence is available at present to warrant
further discussion on the evolutionary aspects of polymorphism in Macropleurodus.
Ecology
Habitat. M. bicolor is widely distributed within Lake Victoria, occurring most
frequently in littoral and sublittoral regions, especially where the bottom is hard
(sand, rock or shingle), but only rarely over mud. Depth distribution is fairly
restricted, with a maximum of between 30 and 40 feet (see also Graham, 1929).
Food. From gut analyses of numerous specimens (throughout the size range
60-150 mm.) it is apparent that snails and insect larvae are the predominant food
organisms (see also Graham, 1929). Shell fragments are rarely found in the stomach
or intestine although opercula are usually present. Aquarium observations confirm
that almost the entire body of the snail is removed from its shell before ingestion
takes place (vide p. 325), although small snails and, on occasion, thin-shelled species,
may be crushed intra-orally before being swallowed. As a result of this feeding
mechanism snail remains are so fragmentary as to preclude accurate identification ;
remains of Gabbia sp. have, however, been positively identified on several occasions.
The insects most commonly recorded from the pabulum of M. bicolor are larvae
of the boring may-fly, Povilla adusta Navas, with other larval Ephemeroptera, and
larval Chironomidae occurring less frequently. The proportion of insect to molluscan
food eaten is difficult to determine and is probably related to local and cyclic abund-
ance of these organisms.
In the light of numerous gut analyses which are now available for this species,
Graham's record (1929) of fish and cichlid eggs from the stomach of M. bicolor requires
some comment. Apart from this record no other instances of piscivorous habits are
known for M. bicolor. Unfortunately, Graham does not give a detailed analysis of
MONOTYPIC GENERA OF CICHLID FISHES IN LAKE VICTORIA 311
the gut contents, particularly of the number of specimens from which his data are
derived. Bearing in mind this limitation, it is suggested that the fish-remains and
eggs were from the stomachs of different individuals and consequent upon brooding
female fishes swallowing their own young, a relatively common occurrence when the
fishes are caught in gill-nets.
Breeding. The exact spawning grounds of M. bicolor are unknown : eggs and
larvae at all stages of development have been found in the mouths of various female
fishes and it is presumed that the species is a mouth-brooder. Brooding females
have been caught in all parts of the species range. The smallest sexually active fish
examined was a female 96 mm. long [ex Kisumu). The habitat of post-larval fishes
is unknown.
Affinities
Unlike the other monotypic genera M. bicolor can apparently be related to an
extant species of Haplochromis. Similarity in skull architecture of M. bicolor and
H. prodromus has already been noted. There are additional similarities in the short
and stoutly constructed jaws of both species, besides a marked resemblance in
general facies. Against these resemblances must be set the very different tooth form
and dental pattern of H. prodromus, although in this species the teeth are stout and
the inner series well developed. The structure of the head and dental patterns
suggest that, at a functional level, the condition represented by H. prodromus might
well be considered pre-adaptive to the development of a relatively massive dentition,
such as that of M. bicolor.
Study material and distribution records
Museum and Reg. No.
S.L. (mm.).
Locality.
Collector.
British Museum (N.H.) :
1906. 5. 30. 414 (type of H. bicolor)
1906.5.30.378 ....
1928.5.24.493-503
1928.5.24. 1-3 ....
125
115
115-125
120-125
Bunjako (Uganda)
Ditto
See below
Ditto
Degen
. M. Graham
Ditto
Paris Museum :
12.278 (holotype of H. mater-
familias .....
12.279-281 (paratypes of H. mater-
familias ; proportions not in-
cluded in description above)
109
91-97
Port Florence,
Kenya
Ditto
. Alluaud and
Jeannel
Ditto
Genoa Museum :
(Type of Bayonia xenodonta Blgr.) .
73
Jinja
Bayon
B.M.(N.H.) :
1955.2. 10. 5-11 ....
78-110
Kisumu
. E.A.F.R.O.
1955.2. 10.50-57
101-123
Jinja, Napoleon
Gulf
Ditto
1955.2. 10.43-46
79-104
. Beach nr. Nasu Point,
Buvuma Channel
•
3 i2 MONOTYPIC GENERA OF CICHLID FISHES IN LAKE VICTORIA
Museum and Reg. No.
S.L. (mm.).
Locality.
Collector.
B.M.(N.H.) :— cont.
1955.2. 10.22-23
82 and 97
Grant Bay (Uganda) .
E.A.F.R.O
1955.2. 10.21
102
Dagusi Island
Ditto
1955.2. 10.65-73
105-120
. Off southern tip of
Buvuma Is.
"
1955.2. 10.24-42 and 74-83 .
90-122
. Harbour at Entebbe .
,,
1955.2. 10.58-62
58-67
Bugonga Beach
,,
1955.2. 10.12, 16-20 and 48 .
112-130
Busongwe Bay
(Kagera River mouth)
>>
1955.2. 10.63-64
. no and 147
Ma jit a Beach
(Tanganyika Terr.)
i >
1955.2. 10.4, 49 and 84
115-119
Mwanza
,,
Graham (1929) records the occurrence of M. bicolor as follows :
Kenya : Kavirondo Gulf : Off Sukuri Island.
Off Ulambwi bay.
Mbita Passage.
Near Nzoia River.
Kadimu Bay.
Tanganyika Territory : Mwanza.
Smith Sound.
Genus PLATYTAENIODUS Boulenger, 1906
Platytaeniodus Boulenger, 1906, Ann. Mag. nat. Hist. (7) 17, 451 ; Idem, 1907, Fish. Nile, 493 ;
Idem, 1915, Cat. Afr. Fish. 3, 426, fig. 292 ; Regan, 1922, Proc. zool. Soc, Lond. 190. Type
species : Platytaeniodus degeni Blgr. 1906.
Generic synonyms :
Astatotilapia (part) "\ ^ , ,
tt u 7 • / 1\ >For references see synonymy under species.
Haplochromis (part) J J J J r
Generic characters and synonomy
Both Boulenger and Regan considered the premaxilla and its tooth pattern
diagnostic. Boulenger (1914) states :
"... the alveolar surface of the premaxilla widening towards the pharynx, the
band of teeth in the upper jaw horseshoe shaped ..."
Additional material shows, however that in fishes below 100 mm. standard length
the posterior premaxillary dentigerous (alveolar) surfaces are not always expanded
medially. Nevertheless, even in small specimens the premaxilla is stouter and its
dentigerous surface wider than in Haplochromis ; posteriorly the teeth are arranged
in several rows, so that the premaxillary tooth band is always clearly U-shaped,
with the arms at least as broad as the medial part. Broadening of the posterior
alveolar surfaces is gradual and shows positive allometry with standard length ; in
some large individuals the left and right surfaces are closely apposed in the mid-line
(Fig. 6, c).
MONOTYPIC GENERA OF CICHLID FISHES IN LAKE VICTORIA 313
Tooth bands in the lower jaw are more readily diagnostic and less subject to
variation with absolute size than those of the premaxilla. In Platytaeniodus the
mandibular teeth are confined to the anterior and antero-lateral portions of the
dentary and are grouped into two broad and roughly pyriform patches, contiguous
at the symphysis ; posteriorly there is a short, single row of four to seven teeth
Icm.
B
Fig. 6. — Platytaeniodus degeni. (a) Premaxilla, (b) dentary, both in occlusal view.
Skeleton prepared from a specimen 80 mm. S.L. (c) Mouth of the holotype, ca. x 4.
[(c) from Boulenger, Fishes of the Nile.']
314 MONOTYPIC GENERA OF CICHLID FISHES IN LAKE VICTORIA
lying between the ascending part of the ramus and the anterior tooth bands (Fig. 6,
b and c).
When provisionally referring Astatotilapia jeanneli Pellegrin to Haplochromis
macrops Blgr., Regan (1922) was apparently misled by the large eye and shallow
preorbital of A. jeanneli. There is undoubtedly some resemblance between H.
macrops and P. degeni, but this is confined to superficial characters, and is belied
by their fundamentally different dentition. Re-examination of the types of A.
jeanneli reveals that the premaxillary teeth are arranged in four or five series of equal
breadth both laterally and posteriorly, a condition never observed in H. macrops.
Furthermore A . jeanneli has the mandibular teeth grouped anteriorly and laterally
in five series, with only a short, single series posteriorly. That is to say, both type
specimens have a dentition typical for small P. degeni. In other characters too,
notably the mouth with its broad lower j aw, shorter than the upper, and the almost
completely hidden maxilla, A . jeanneli agrees more closely with P. degeni than does
H. macrops or any other Haplochromis species.
On these grounds, therefore, I consider A. jeanneli to be synonomous with P.
Diagnosis
Cichlid fishes of the Haplochromis group, but differing from Haplochromis in
having broad bands of teeth on the posterior part of the premaxillary dentigerous
surface, which is expanded medially in large specimens but is of almost equal breadth
anteriorly and laterally in fishes of less than 100 mm. S.L. Teeth on the dentary
grouped into two, broad, pyriform, curved and contiguous bands anteriorly and
antero-laterally, but continued posteriorly as a short single row only. Lower jaw
usually shorter than the upper ; maxilla almost completely hidden below the
preorbital.
Fig. 7. — Platytaeniodus degeni, £. Drawn by Miss L. Buswell,
MONOTYPIC GENERA OF CICHLID FISHES IN LAKE VICTORIA 315
Platytaeniodus degeni Boulenger, 1906
(Fig. 7)
Platytaeniodus degeni Boulenger, 1906, I.e.; 1907, I.e. and pi. xci, fig. 1; 1915, I.e. and fig. 292;
Regan, 1922, 190, fig. 14.
Astatotilapia jeanneli Pellegrin, 1913, Bull. Soc. zool. France, 37, 313.
Haplochromis jeannelli (Pellegrin), Blgr. 1915, Cat. Afr. Fish. 3, 291.
Haplochromis maerops (part) Regan, 1922, Proc. zool. Soc. London, 166.
Description
Thirty-six specimens (size range 67 to 154 mm.) comprising the type and other
specimens in the British Museum (Nat. Hist.) (including material newly collected
by E.A.F.R.O.), as well as the types of A. jeanneli are considered in this description.
Since most characters tabulated below show some allometry with standard length
these data are grouped into two size classes. In some characters intra-group vari-
ability is high, but further subdivision into smaller groups is impracticable.
In the table of proportions, head length, depth of body and length of caudal
peduncle are expressed as percentages of standard length ; all other characters are
expressed as percentages of head length. Range and mean are given for each
character.
Table I.
Standard Length.
Depth of body .
Length of head
Depth of preorbital .
Least interorbital width
Length of snout
Diameter of eye
Depth of cheek
Length of lower jaw .
Caudal peduncle
67-93 mm.
(23 specimens)
32-5-40-5
M =35-6
30-0-36-5
M = 32-4
12-0-16-0
M = 14-2
25-0-30-8
M = 27-1
25-0-32-0
M = 30-0
27-4-33-4
M = 30-8
18-5-26-1
M = 19-7
32-1-39-6
M = 37-0
13-9-21-5
M = 16-5
98-154 mm.
(13 specimens).
34.78-43-0
M = 38-4
31-5-34-7
M = 32-7
13-7-17-4
M = 15-2
28-2-34-8
M = 31-2
31*75-37-4
M = 34-8
22-1-26-0
M = 24-8
20-0-26-6
M = 23-4
35-0-38-1
M = 36-0
14-0-18-5
M = 16-7
Dorsal profile of head and snout gently (in a single specimen somewhat strongly)
decurved ; mouth horizontal ; lower jaw equal to or more usually shorter than upper,
its length /breadth ratio i-i6-i-68 : in some specimens the lower jaw, including
lips, is slightly broader than the upper. Lips well developed and somewhat thickened ;
maxilla almost completely hidden beneath the preorbital, with only its postero-
ventral tip exposed and extending to below the anterior orbital margin, or slightly
beyond.
3 i6 MONOTYPIC GENERA OF CICHLID FISHES IN LAKE VICTORIA
The holotype, a male of 115-0 mm. S.L., is figured by Boulenger (1907 and 1915).
This fish is both somewhat atypical and slightly distorted in preservation and the
impression given of a deep ventral profile and a slightly oblique mouth is not
characteristic.
Gill rakers. Short and stout, 7-8 on lower limb of the anterior arch, the lowest one
or two usually reduced.
Dentition. Premaxilla with 4-8 rows of teeth ; dentary with 4-6 rows.
With few exceptions, specimens over 100 mm. S.L. have the posterior part of the
premaxillary dentigerous surface expanded medially, so that the upper tooth bands
are broader posteriorly than anteriorly. In small individuals, although there are
several series of teeth posteriorly, the tooth band is either of equal width at all
points, or medial expansion of the premaxilla may have begun, causing the band to
be very slightly wider posteriorly (Fig. 6, a). Expansion of the posterior surface is
not correlated with an increase in the number of tooth rows borne on it, which are,
in fact, equal to or slightly fewer than those on the anterior part of the premaxilla.
Mandibular dental pattern as described for the genus ; pyriform band from half
to two thirds as broad as long.
Teeth are variable in form, those of the outer series slenderly conical, with or
without an admixture of unequally bicuspid teeth. Teeth of the inner series all
unicuspid in most specimens above 100 mm. S.L. and in some below that size ;
otherwise the outermost teeth unicuspid and the remainder tricuspid or exceptionally,
bicuspid.
Scales ctenoid ; lateral line interrupted, with 31 (f.2), 32 (f.5), 33 (f.14), 34 (f.io),
35 (f.2), or 36 (f.3) scales. Cheek with 3-4 series of imbricating scales ; 7-9 scales
between lateral line and origin of dorsal fin ; 7-9 (rarely 10) between the pectoral
and pelvic fins.
Fins. Dorsal with 24 (f.3), 25 (f.23) or 26 (f.io) rays, anal with 11 (f.6), 12 (f.23)
or 13 (f.7) rays, comprising XV-XVII 8-10 and III 8-10 spinous and soft rays.
Pectoral fin 22-6-31-8 (M = 28-0) per cent of standard length. Caudal truncate,
scaled on its proximal half only. Pelvics with the first ray produced, extending to
the vent in a few specimens and to the spinous part of the anal in most.
Syncranium. Since the form of premaxilla and dentary in P. degeni is correlated
with the well-developed tooth pattern, both these bones depart very strikingly from
the typical Haplochromis condition.
In small individuals of P. degeni, the premaxilla bears a superficial resemblance
to that of Hoplotilapia, particularly with regard to the dental pattern, but in large
fishes it is unique. On the other hand, the dental pattern and morphology of the
dentary are comparable in both large and small individuals. The dentary is charac-
terized by its broad and laterally expanded anterior tooth-bearing portion, which
imparts to this bone an appearance unique amongst the Lake Victoria cichlids.
The neurocranium of P. degeni is intermediate between the generalized Haplo-
chromis type and that of M. bicolor. It is strictly comparable with the neurocrania
of species of the H. crassilabris group. Here the skull is characterized by a somewhat
shortened and steeply inclined ethmo-vomer complex ; in consequence, the anterior
skull profile is also steep. In a typical Haplochromis skull the ethmo-vomer is longer
MONOTYPIC GENERA OF CICHLID FISHES IN LAKE VICTORIA 317
and rises less steeply, meeting the downward sloping frontals at a wide angle. As a
result, the anterior profile is shallower and also more acute than in the " crassilabris "
type skull (Fig. 5).
Jaw musculature in P. degeni dos not differ greatly from that of a generalized
Haplochromis species.
Coloration. Preserved material. Males : dusky to dark grey ; dorsal and anal
fins dark ; pel vies black, caudal colourless. Females and immature individuals :
silver-grey or light brown ; fins colourless.
Transverse and longitudinal banding sometimes occurs, being most clearly marked
in females and young individuals ; when present, there is a well marked median
longitudinal stripe, a fainter and interrupted band running slightly below the dorsal
fin, and eight or nine narrow transverse stripes on the flank and caudal peduncle.
Faint lachrymal and interocular stripes may also be present. The presence and
intensity of these markings is apparently related to the emotional state of the fish
or may only appear after death.
Coloration in life. Sexually active males : ground colour light blue-grey, lips
iridescent blue. Chest and branchiostegal membrane black. Fins ; dorsal sooty,
lappets and spots on soft part red ; caudal with red flush, most intense along margin ;
anal with dusky pink flush and several yellow ocelli. Females : ground colour
golden fawn ; all fins neutral, dorsal with orange lappets and spots, especially on
the soft part ; caudal with orange margin and maculae ; anal with faint or well
marked yellow ocelli.
Ecology
P. degeni is recorded from several areas in Lake Victoria (see below), but as so
few specimens are known it is not possible to generalize on habitat preferences.
From the scanty data available it appears that the species is probably restricted to
littoral and sub-littoral regions where the water is less than fifty feet deep. Specimens
have been caught in nets set over both hard and soft substrates, but the greater
number came from stations having a ! sand or shingle bottom.
Food. The distinctive dentition of this species suggests a highly specialized diet.
Tantalizingly few fish, however, have yielded ingested material. Twelve specimens
have been examined, all of which were caught in nets set overnight or in seines
operated during varied daylight hours. According to the substrate over which they
were living, ten fishes had either sand grains or organic mud in the stomach and
intestines, together with fairly dense aggregations of mucus. Two fishes, caught on
different occasions at a station near the southern tip of Ramafuta Island (Buvuma
Channel), had the entire alimentary tract filled with the diatom Melosira. Diatoms
from the stomachs of these fishes showed only slight signs of digestion, but samples
taken from the mid-intestines and recta were almost completely digested. Animal
remains, occurring sporadically, included insect larvae, Hydracarina, fragments of
Copepoda, Ostracoda, and in two specimens shell fragments of Pelecypoda (Sphae-
riidae).
Most guts also contained some diatoms and blue-green algae, the former apparently
digested, the latter intact. The very small quantity of ingested material in any one
318 MONOTYPIC GENERA OF CICHLID FISHES IN LAKE VICTORIA
individual is striking and no particular organism, or group of organisms, occurs with
sufficient frequency to indicate what the food of P. degeni may be. Since sand and
bottom debris is significant in the majority of specimens, it is possible that the species
may feed on the micro-fauna and flora living on and within the substrate. Thus,
broad bands of jaw teeth may serve to rasp and loosen food from the surrounding
sand.
Breeding. Spawning sites are unknown ; only two females, both from beaches in
the Mwanza area, have been found with eggs in the mouth. The smallest individual
with demonstrably active gonads was a female, 71-0 mm. long.
Affinities
There is no obvious relationship between P. degeni and any known Lake Victoria
species or species-group of Haplochromis, with which genus the species shows
fundamental affinities. The peculiar premaxillary and mandibular tooth patterns
serve to set P. degeni apart from even those Haplochromis with several series of
inner teeth. Regan (1922) considered P. degeni as being " very near " to H. prod-
romus, which species he believed " shows a slight departure from the normal Haplo-
chromis dentition towards the Platytaeniodus type ". His opinion was based on the
holotype and then unique specimen of H. prodromus. Summarizing unpublished
data on H. prodromus, it is clear that the type specimen has an aberrant dental
pattern and that its resemblance to P. degeni is purely superficial. Whereas in large
Platytaeniodus there is an actual expansion of tooth-bearing surfaces, in H. prodromus
only the tooth-band is apparently expanded ; its increased breadth is actually due
to the posterior teeth being more widely separated from one another than are the
anterior teeth. In no specimen of H. prodromus is the upper tooth band as broad
posteriorly as anteriorly, yet this is the usual condition in P. degeni. Further, the
dentary of P. degeni differs considerably from that of H. prodromus. The evolution
of a wholly multi-seriate dentition has probably occurred more than once within the
Lake Victoria species-flock, as for instance in the H. sauvagei group and again in
the monotypic genera. Thus any apparent relationship between P. degeni and H.
prodromus should be considered as consequent upon convergent evolutionary trends,
the ultimate expressions of which are achieved by manifestly dissimilar means.
Study material and distribution records
Museum and Reg. No.
S.L. (mm.).
Locality.
Collector.
British Museum (Nat. Hist.) :
1906. 5. 30. 51 1 (holotype)
114
. Bunjako (Uganda)
Degen
1909-3-29.10
98
Sesse Is. (Uganda)
1928.5.24 ....
93
Mbita Passage
(Kenya)
. M. Graham
Paris Museum :
12 . 262 (holotype of A . jeanneli)
72
Port Florence
. Alluaud and
(Kenya)
Jeannel
12 . 262 (paratype of A . jeanneli)
67
Ditto
Ditto
MONOTYPIC GENERA OF CICHLID FISHES IN LAKE VICTORIA 319
Museum and Reg. No. S.L. (mm.). ' Locality. Collector.
B.M. (N.H.) :
1955.2. 10.91-94 .... 73-79 • Kisumu (Kenya) . E.A.F.R.O.
1955-2.10.105-106 . . . 78-90 . Kamarenga (Kenya) . Ditto
1955-2. 10. 115 .... 74 Kendu (Kenya)
1955.2. 10.88 .... 74 Likungu (Kenya) . „
Genus HOPLOTILAPIA Hilgendorf, 1888
Hoplotilapia Hilgendorf, 1888, S.B. Ges. naturf. Fr. Berlin, 76-77 (type species (Paratilapia ?)
retrodens Hilgendorf, I.e.) ; Regan, 1922, Proc. zool. Soc, Lond. 190.
Cnestrostoma Regan, 1920, Ann. Mag. nat. Hist. (9) 5, footnote p. 45 (type species Paratilapia
polyodon, Blgr.).
Haplochromis (part) "]
Paratilapia (part) }■ For references see synonymy under species below.
Hemichromis (part) J
Generic characters and synonymy
The holotype of Hoplotilapia retrodens (in the collections of the Zoologisches
Museum der Humboldt-Universitat, Berlin) has not been examined by me, nor can it
be definitely established whether this specimen is still in existence. Dr. Kurt Deckert
of the Zoologisches Museum has, on two occasions, kindly attempted to locate
several specimens, including the type of H. retrodens. Of these he writes : " Ich
muss Ihnen leider mittheilen, dass unser Suchen nach den verlangten Typen ohne
Erfolg geblieben ist, obwohl ich mit grosser Sicherheit anneheme, dass sie nicht
verloren-gegangen sind. ' '
Hilgendorf s original description of {Paratilapia?) retrodens (1888), although brief
and lacking detail, nevertheless stresses characters which clearly separate this
species from others of the Haplochromis group : viz. a multi-seriate dental pattern
with stout and enlarged posterior teeth. Pfeffer's redescription (1896) of the same
specimen confirms and extends this account. In the material at my disposal, how-
ever, the posterior teeth are clearly enlarged only in the upper jaw, and not in both,
as stated for the type. Specimens described below agree closely with the type in
the other characters described by Hilgendorf and Pfeffer ; slight differences in scale
numbers can probably be attributed to different methods of making these counts.
In addition to the four specimens of H. retrodens in the British Museum (Nat.
Hist.) it has been possible, through the kindness of Dr. Delfa Guiglia, to examine
the type of Paratilapia polyodon Blgr. and one other specimen (Museo Civico
di Storia Naturale, Genoa, reg. no. G.E. 12.994) determined by Boulenger as
P. polyodon, and to confirm Regan's view (1922) that these are conspecific with
H. retrodens.
Diagnosis
Differing from Haplochromis as defined by Regan (1920 and 19226) in having
broad bands of teeth in both jaws, well developed and usually of almost uniform
breadth throughout or very slightly narrower posteriorly. Posterior teeth of the
upper jaw enlarged and stout, those of the lower jaw slightly, if at all, enlarged, but
320 MONOTYPIC GENERA OF CICHLID FISHES IN LAKE VICTORIA
the tooth-band continued posteriorly on to the steep ascending contour of the
dentary. Lower jaw wide and flat, almost square in anterior outline, slightly shorter
than the upper.
Since only three of the collected specimens of this genus are small it is not possible
to generalize on differences which apparently exist between the dentition of adult
Icm.
Fig. 8. — Hoplotilapia retrodens. (a) Neurocranium, (b) dentary in lateral view,
(c) dentary, occlusal view. Dentition only part indicated. From a specimen of 125
mm. S.L.
MONOTYPIC GENERA OF CICHLID FISHES IN LAKE VICTORIA 321
and juvenile fishes. In these three specimens (74-0, 76-0 and 55-0 mm. S.L.) the
tooth bands are broad anteriorly, being composed of 5, 3, and 3 series respectively.
Laterally, however, they are reduced to two series, whilst postero-laterally only the
outer series persists. In none is the dentition continued onto the ascending part of
the dentary, although the shape of the lower jaw is as in the adult.
Two adults of 96 and no mm. S.L., collected in a single seine haul at Bukakata,
retain these presumedly juvenile dental characters. They form a graded morpho-
logical series with a third specimen (134-0 mm. S.L., from the same station) which
exhibits only slight departure from the " typical " condition.
The broad and shallow lower jaw (Fig. 8, Band c) of typical individuals is unique
amongst Lake Victoria Cichlidae. A few specimens of H. retrodens have the dentary,
at least in external appearance, similar to that of Haplochromis, although in every
case the dental pattern is typical for Hoplotilapia.
Fig. 9. — Hoplotilapia retrodens, $. Drawn by Miss L. Buswell.
Hoplotilapia retrodens Hilgendorf, 1888
Fig. 9.
(Paratilapia ?) retrodens Hilgendorf, 1888, S.B. Ges. naturf. Fr. Berlin, 76.
Hoplotilapia retrodens Hilgendorf, t.c, 77.
Hemichromis retrodens, Pfeffer, 1896, Thierw. O. Afr. Fische, 19.
Haplochromis bicolor (part) Boulenger, 1906, Ann. Mag. nat. Hist. (7) 17, 444 ; two specimens
B.M. no. 1906. 5. 30. 417 and 418.
Paratilapia bicolor (part) Boulenger 1915, Cat. Afr. Fish. 3, 346.
Paratilapia poly don Boulenger, 1909, Ann. Mus. Genova (3) 4, 306, rig. ; Idem, 191 1, ibid. 5,
68 ; Idem 1915, Cat. Afr. Fish. 3, 349, fig. 236.
Cnestrostoma polyodon (Boulenger), Regan, 1920, Ann. Mag. nat. Hist. (9) 5, footnote p. 45.
322 MONOTYPIC GENERA OF CICHLID FISHES IN LAKE VICTORIA
Description
From available material it would appear that only the interorbital width shows
marked allometry with standard length. It must, however, be borne in mind that
paucity of specimens within the smaller and larger size groups may obscure such
relationships, especialy since, in those size-groups which are well represented,
individual variability is high. With the exception therefore of interorbital width,
ranges and means are given for the sample as a whole. For interorbital width the
range and mean are given for three size groups, 74-115 mm. (N = 21), 116-130 mm.
(N = 32), and 131-144 mm. (N = 11).
Depth of body 31-6-41-6 (M = 38-3), length of head 30-1-34-8 (M = 32-0) per
cent of standard length. Dorsal profile of head and snout straight or slightly concave,
steeply sloping. Preorbital depth 12-5-18-8 (M = 16-4) per cent of head length ;
least width of interorbital 24-2-33-8 (M = 28-7), 28-6-35-1 (M = 31-5) and 31-1-
33-4 (M = 31*6) per cent for the three size groups respectively ; length of snout
29-0-36-8 (M = 32-5), eye 23-8-29-6 (M = 26-9), depth of cheek 19-5-28-2 (M =
23-9) per cent of head length.
Lower jaw slightly shorter than upper 33-7-40-8 (M = 38-2) per cent of head, the
length /breadth ratio from broader than long to 1-33 times as long as broad. Mouth
horizontal, lips somewhat thickened. Posterior tip of the maxilla extending to the
vertical from the anterior margin of the orbit or as far as the pupil.
Described from 64 specimens, 55-144 mm. standard length.
With few exceptions, there is remarkable uniformity in the general facies of H.
retrodens. In this respect the figure of Paratilapia polyodon type specimen in
Boulenger (1909 and 1915) can be considered fairly representative. The lower jaw
in this specimen has, however, been broken and subsequently distorted in preserva-
tion, consequently the ventral head profile of the figured specimen is inclined upward
and is not horizontal, as it would be in life. The greatest departure from typical
physiognomy and body form is seen in a single specimen from Bukoba (Tanganyika)
which has the body relatively elongate and the head profile strongly curved. Despite
this aberrance in gross morphology, the dentition and other fundamental characters
of this specimen are typical.
Teeth and dental pattern. Fishes above 90 mm. S.L. have 5 to 8 series of teeth
anteriorly and 4 to 5 series posteriorly in the upper jaw ; 5 to 8 (rarely 9 or 10)
series anteriorly and 3 to 5 series posteriorly in the lower jaw. From 40 to 68 teeth
in outer series of the upper jaw. Teeth small, those of the outermost series largest,
variable in form, but usually unicuspid ; some bi- and tricuspid teeth occur in the
inner series. Two specimens from Majita and Mwanza (Tanganyika Territory) have
markedly aberrant, stoutly conical or nearly molariform teeth in all series of both
jaws. In both specimens the dental pattern is otherwise typical.
Syncranium. The premaxilla is comparable in form with that of Haplochromis,
except that it is stouter, especially posteriorly, and its dentigerous surface broader.
As noted above, the broad and greatly flattened dentary is unique amongst Lake
Victoria cichlids (Fig. 8).
The neurocranium agrees closely with that of Platytaeniodus and with Haplo-
chromis of the " crassilabris " species-group (see p. 316).
MONOTYPIC GENERA OF CICHLID FISHES IN LAKE VICTORIA 323
Pharyngeal bone with a triangular and approximately equilateral dentigerous
surface ; pharyngeal teeth cuspidate, the median series often enlarged.
Scales ctenoid, lateral line interrupted, with 31 (f.i), 32 (f.io), 33 (f.38), 34 (f.14)
or 35 (f.i) scales. (Hilgendorf (1888) gives L.S. 30 [probably on a mid-lateral series]
for the holotype.) Cheek with three or four series of imbricating scales. (Hilgendorf
[loc. cit.) gives 4-5 rows) ; y-S scales — rarely 6 — between lateral line and origin of
dorsal fin ; 8-9, less commonly 7, between pectoral and pelvic fin bases.
Fins. Dorsal with 24 (f.4), 25 (f.45) or 26 (f.13) rays, comprising XV-XVII 8-10
spinous and soft rays. Two specimens have the formulae XIV 8 and XVII 6, but
there are indications that these are the results of wounding and subsequent irregular
healing. Anal with n (f.6), 12 (f.54) or 13 (f.4) rays, i.e. Ill, 8-10. Caudal fin trun-
cate, scaled on proximal half only. Pectoral fin 23-9-33-0 (M = 28-4) per cent of
standard length. Pelvics with first ray produced in both sexes but proportionately
longer in adult males.
Coloration. Colours of preserved specimens are variable. Adult males : dark
grey to black, median fins dark, caudal colourless or maculate ; pelvics black.
Females and immature males : light olive brown to silver, all fins yellow or colour-
less. Sometimes faint traces of transverse bars, an interrupted band below insertion
of dorsal fin and a well-marked mid-lateral stripe.
Colours in life. Adult males : ground colour dark grey-green (darkening to deep
slate-grey immediately after death), chest and branchiostegal membrane black.
Dorsal fin dark grey-green, lappets red ; red spots, often coalescing into streaks,
between the rays. Caudal with proximal third to half black, distal part red. Anal
with dark base, remainder red ; ocelli yellow. Pectoral colourless ; pelvics black,
with faint red flush along median rays. Adult females : ground colour light olivaceous
dorsally, shading through sulphur-yellow to pearly-white ventrally ; usually two
longitudinal bands as described above, of variable intensity. Lips yellow. Dorsal
and caudal fins olivaceous ; anal colourless or with slight yellow flush ; small and
ill-defined orange spots sometimes occur in the position of the male ocelli. Pelvics
and pectoral fins yellow.
Three specimens (2 ?, 1 <J) from Bukakata, which were noted when discussing
atypical dental patterns (p. 321), also exhibit aberrant coloration. This is known
only from preserved material, in which it has the form of three large and elongate
black spots, arranged mid-laterally, on a light ground colour.
Almost completely sex-limited polychromatism occurs in this species and, as in
M. bicolor, it is the female which usually exhibits atypical coloration. Besides the
black and yellow piebald, as described for M. bicolor, there is a second and more
colourful, if less distinctive, pattern. Any attempt to describe this pattern must
perforce be imprecise, since intensity and detail show a remarkable range of indi-
vidual variability. The ground colour is invariably a light sandy-yellow, with a
superimposed orange flush, usually most intense on the head. Dorsally there are a
number of irregular and ill-defined dark blotches, separate or confluent, which occa-
sionally extend on to the flank. In some individuals this dorsal pigmentation is
comparable with the clearly defined blotches of a typical bicolor pattern ; in others
it is more diffuse and individual elements are only faintly discernible. The whole
324 MONOTYPIC GENERA OF CICHLID FISHES IN LAKE VICTORIA
body, including the fins, is also peppered with small melanophores, particularly
on the head and opercula. Fin coloration is extremely variable, but the caudal and
anal are usually flushed with bright orange.
Sampling bias in favour of atypically-coloured individuals undoubtedly occurs
and precludes the accurate estimation of frequency. Of thirty-five females examined,
thirteen were " bicolor " and eleven of the other pattern. Polychromatism is known
in populations from most areas in which the species has been collected.
A single male (119-0 mm. S.L., Busongwe, Kagera River area) had an incipient
bicolor pattern, resembling in appearance and degree of pigmentation one of the
" bicolor " males of M. bicolor (p. 309).
The genetical basis of, and selection factors maintaining polychromatism in H.
retrodens remain undetermined, as previously explained in the case of M . bicolor.
Ecology
H. retrodens is widespread throughout Lake Victoria (see below). Sampling in
many habitats shows the species to be restricted to littoral and sub-littoral areas,
especially where the substrate is hard, and usually where there is submerged vegeta-
tion. The majority of specimens was caught in water from twelve to twenty feet
deep, with a few from slightly deeper water.
Food. The gut contents of sixty-five individuals (size-range 75-0-144-0 mm. S.L.)
have been examined ; of these, only seven were empty. Mollusca (particularly
Lamellibranchiata) are the predominating food, being recorded from thirty-six
fishes. The majority of Mollusca is represented by finety broken shells. Insecta
(f.8) — especially Ephemeroptera, Trichoptera and Chironomidae — together with
Crustacea (f.3) and Hydracarina (f.2) occur less frequently. In fifteen fishes the
ingested material consisted almost entirely of fragmented plant epidermis, whilst
eleven others contained only sand grains or organic mud.
Observations made on fishes living in aquaria show that H. retrodens, when feeding,
repeatedly makes short, darting movements into or over the substrate, the broad
and horizontal mouth serving as a scoop or shovel. Much bottom material is spilled
from the mouth or with the exhalent current. Snails were retained and apparently
crushed within the jaws before being passed back to the pharynx. The presence of
macerated plant epidermis in several fishes may be explained either as ingested
bottom debris, or as the result of the fish actively scraping the leaves and stems of
submerged plants, for which purpose the multi-seriate dentition would seem adapted.
It may be significant to note that the frustules of epiphytic diatoms found in the
stomachs of these fishes were empty, as were the epidermal cells which had been
ruptured.
Breeding. No information is available on the breeding habits of H. retrodens. The
smallest sexually mature individual was a female of 96 mm. S.L. (Bukakata, Uganda).
Affinities
Morphologically, Hoplotilapia retrodens, like P. degeni is relatively far removed
from any Lake Victoria Haplochromis species or species group. Neither is it closely
MONOTYPIC GENERA OF CICHLID FISHES IN LAKE VICTORIA 325
related to the other monotypic genera of that lake. Regan's remark (1922, p. 159)
that " A remarkable group of three species includes Haplochromis sauvagei and the
monotypic genera Macropleurodus and Hoplotilapia, which scarcely differ from each
other except for the considerable differences in dentition, ... "is difficult to
endorse, unless the various structures whose forms are modified in association with
the dentition and feeding habits are included in the term " dentition ". Macro-
pleurodus and Hoplotilapia have manifestly dissimilar jaw morphology and also
differ in the form of the neurocranium. These structural differences impart a charac-
teristic physiognomy to the species, which allows them to be recognized without
reference to the dental form and pattern. The occurrence of " bicolor " patterns in
both M. bicolor and H. retrodens is suggestive, but at present little importance can
be attached to this character since it occurs in at least three widely divergent species
of Haplochromis as well as in M. bicolor. If neurocranial form can be considered as
being of phylogenetic value, then the affinities of Hoplotilapia lie with the H. crassi-
labris species group, although H. retrodens has departed considerably from this
complex in the form of its jaws and dental pattern.
Hoplotilapia and Macropleurodus show interesting trophic parallels, both between
the two genera and in relation to the three mollusc-eating cichlids of Lake Victoria,
viz. Astatoreochromis alluaudi, Haplochromis pharyngomylus and H. ishmaeli.
Ecologically, Hoplotilapia and Macropleurodus occur together with A. alluaudi and
H. pharyngomylus, but the diet of Hoplotilapia and especially of Macropleurodus,
unlike that of the two last-named species, includes a substantial number of insects.
The parallelism between Hoplotilapia and Macropleurodus extends to the method
of feeding, and particularly the manner in which the hard-shelled prey is crushed. It
is this characteristic which most clearly emphasizes the morphological disparity
between the two genera under discussion on the one hand and the three mollusc-
eating species on the other ; these latter species, although including a monotypic
genus, have deviated less markedly from the generalized Haplochromis anatomy.
Whereas in Astatoreochromis alluaudi, H. ishmaeli and H. pharyngomylus the food
is crushed entirely by means of the hypertrophied pharyngeal bones and teeth, in
Hoplotilapia and Macropleurodus the food is broken mainly by the peculiarly
developed jaws and oral dentition, although the relatively poorly-developed
pharyngeal mill doubtless continues the process.
Despite functional similarity in the jaws and dentition of Hoplotilapia and
Macropleurodus, there is considerable divergence in the detailed morphology of these
elements. On the one hand, in Macropleurodus the jaws are short and stout, with a
narrow gape ; associated with the stout supporting skeleton, the teeth are strong.
On the other hand, the dentary and premaxilla of Hoplotilapia, although broad and
encompassing a wide gape, appear relatively fragile. The dentition of Hoplotilapia,
when compared with Macropleurodus, is seen to be composed of small and slender
teeth which are arranged in bands broader both anteriorly and posteriorly than the
corresponding teeth of Macropleurodus. Since the shells of Gastropoda in Lake
Victoria are stouter than those of Lamellibranchiata, it would not be unreasonable
to suppose that the strong, laterally concentrated and enlarged teeth of Macro-
pleurodus, and the uniformly multi-seriate, finely-pointed teeth of Hoplotilapia, are
326 MONOTYPIC GENERA OF CICHLlD FISHES IN LAKE VICTORIA
associated with the predominantly gastropod diet of the former species, and the
predominantly lamellibranch diet of the latter.
Study material and distribution records
Museum and Reg. No.
S.L. (mm.)
Locality.
Collector.
British Museum (Nat. Hist.) :
1906. 5. 30. 417-418
. 90 and 106
. Buganga (Uganda) .
Degen
1909.5.4. 16
112
Sesse Is. (Uganda)
Bayon
1911.3.3.34
144
Jinja, Ripon falls
(Uganda)
"
1928.5.24.489-492
125-132
Lake Victoria
Graham
Genoa Museum :
C.E. 12.995 (holotype of P. polyodon) 135
—
—
C.E.12.994 ....
112
—
—
B.M. (N.H.) :
1955.2. 10. 141, 147-149.
128-142
. Rusinga Is. (Kenya) .
E.A.F.R.O
1955.2. 10. 145
129
. Homa Bay (Kenya) .
Ditto
1955.2. 10. 142-144, 146.
105-135
. Kamaringa (Kenya) .
,,
1955.2. 10. 137-140
104-117
Kisumu (Kenya)
,,
1955.2.10.116-123,180
76-138
Jinja (Uganda)
,,
1955.2. 10. 177-178
55
. Beach nr. Nasu Point.
,,
(Buvuma Channel, Uganda)
1955.2. 10. 170-171
. 105 and 107
Pilkington Bay
(Uganda)
"
1955.2. 10. 176
114
Ramafuta Is.
(Uganda)
"
1955.2. 10. 168-169
155
. Yempita Is. (Rose- .
bery Channel, Uganda)
y >
1955.2. 10. 124-132
117-124
Harbour, Entebbe
,,
1955.2. 10. 133
109
Bugonga beach,
Entebbe
"
1955.2. 10. 172-175
96-142
Old Bukakata
(Uganda)
ll
1955.2. 10. 150-163
104-132
. Busongwe (Kagera .
R. mouth, Uganda)
"
1955.2. 10. 179
127
Beach south of
Bukoba (Tanganyika)
"
1955.2. 10. 164-167
106-117
Majita beach
(Tanganyika)
"
1955.2. 10. 134
86
Harbour, Mwanza
(Tanganyika)
"
1955.2. 10. 135-136
. 74 and 139
. Capri Bay, Mwanza .
(Tanganyika)
"
Graham (1929) lists the distribution of H. retrodens as follows :
Kenya Colony : Mbita passage ; Kavirondo Gulf ; Kadimu Bay.
Tanganyika Territory : Mussonya Bay (Ukerewe Is.); trawl near Bukoba.
MONOTYPIC GENERA OF CICHLID FISHES IN LAKE VICTORIA 327
PARALABIDOCHROMIS gen. nov.
Diagnosis
Cichlid fishes of the Haplochromis group, but differing from that genus in having
the anterior teeth in both jaws procurrent and disproportionately longer than the
adjacent lateral teeth. Jaws narrowing at the symphysis ; lips thickened. Known
only from Lake Victoria.
Type species : Paralabidochromis victor iae sp. nov.
The single specimen of Paralabidochromis available provides an interesting taxo-
nomic and phylogenetic problem. No characters have been found which will dis-
tinguish this fish generically from specimens of the genus Labidochromis Trewavas ;
a genus otherwise known only from Lake Nyasa. Unfortunately comparisons must
be limited to characters apparent in preserved material and then only to the few
specimens available. Nothing is known of the coloration in life of adult males in
either genus. This is regrettable since coloration might well provide a reliable
indication of the affinities of the two genera, both in relation to one another, and to
the species flocks of Victoria and Nyasa (vide Regan, 1921, 686). The presence of a
dark sub-marginal band on the dorsal fin of Labidochromis vellicans, in contradistinc-
tion to its absence in Paralabidochromis is probably of some importance. A sub-
marginal band is not known in any Lake Victoria Haplochromis species, but is
present in most species of the group of Nyasa genera to which L. vellicans is
apparently related (vide Trewavas, 1935, p. 71).
Although on purely morphological grounds it might seem advisable to include
the Lake Victoria species within the genus Labidochromis, such a decision would
imply phyletic relationships between the Victoria and Nyasa species closer than
those between either species and others of its own lake. To avoid this I have given
greater weight to the difference in colour-pattern than would perhaps have been
justified if both inhabited the same lake.
Apart from the presence of the pan- African genera Tilapia and Haplochromis,
there is no obvious relationship between the Cichlidae of Lakes Nyasa and Victoria.
Superficial resemblances between individual species, or genera, in the two lakes have
been associated with differences which point to their being examples of convergent
evolution (Regan, 1922, p. 159), although it would perhaps be preferable to consider
this convergence of morphological characters as parallel evolution since the phenome-
non occurs between species within a group of related genera.
It would seem most probable, therefore, that Paralabidochromis represents a
remarkable example of exact and detailed parallel evolution with Labidochromis.
Apart from the enlarged anterior teeth, neither Labidochromis nor Paralabidochromis
departs greatly from the generalized Haplochromis type, as represented in the rivers
of East and Central Africa. Thus, it is possible that the two genera were independ-
ently evolved from different parental Haplochromis species, which, however, shared
the generalized facies of fluviatile species.
328 MONOTYPIC GENERA OF CICHLID FISHES IN LAKE VICTORIA
Paralabidochromis victoriae sp. nov.
Fig. io.
Description
Depth of body 33-0 ; length of head 31-6 per cent of standard length. Dorsal
profile of head and snout slightly curved and sloping moderately steeply. Preorbital
depth 167 per cent of head length ; interorbital width 25-0, snout length 33-4,
diameter of eye 29-2 and depth of cheek 20-8 per cent of head length. Caudal
peduncle 1-33 times as long as deep, its length 15-8 per cent of standard length.
I cm.
Fig. 10. — Paralabidochromis victoriae, <$, holotype. Drawn by Miss L. Buswell.
Mouth almost horizontal, lips well developed. Posterior tip of the premaxilla
extending to a point nearer the nostril than the anterior orbital margin. The angle
between the rami of each jaw is acute giving a beak-like appearance, which is
enhanced by the peculiar dentition.
Dentition. Thirty-eight teeth in the outer series of the premaxilla ; the six anterior
teeth in both jaws procurrent, greatly elongate, slender, slightly recurved and dagger-
like ; movable (Fig. 10). Postero-lateral outer teeth small and unicuspid, becoming
progressively larger and weakly bicuspid laterally. Inner series in both jaws com-
posed of tricuspid and compressed teeth, of which there are three rows in the upper
and four in the lower jaw ; outermost row of inner teeth in both jaws, somewhat
enlarged and standing slightly apart from the remaining rows.
Lower pharyngeal bone triangular, the teeth small and cuspidate, with only the
median series slightly enlarged.
MONOTYPIC GENERA OF CICHLID FISHES IN LAKE VICTORIA 329
Gill rakers short ; seven on lower limb of first arch.
Scales ctenoid ; lateral line interrupted, with 32 scales. Cheek with 2 scries of
imbricating scales ; 6 scales between origin of the dorsal fin and the upper lateral
line ; 7 between pectoral and pelvic fins. Nuchal scales not exceptionally small.
Fins. Dorsal, XV 8 ; anal, III 9. Pectoral fin 25-0% of standard length ; pelvics
with the first soft ray produced and extending posteriorly to the third anal spine.
Caudal truncate, scaled on proximal half only.
Coloration. Preserved specimen light brown, with indications of seven faint
transverse bands on the flanks, and a dark lachrymal stripe. Pelvic fins black, all
other fins colourless.
Type locality. Sandy littoral, near Nasu Point, Buvuma Channel.
Described from a single specimen, a male 76-0 mm. S.L. (91-0 mm. T.L.), collected
by the author whilst seine-netting at night, 29th May, 1951.
Affinities
Paralabidochromis victoriae is closely related morphologically, to Labidochromis
vellicans of Lake Nyasa, from which it differs principally in possessing large scales
on the chest, cheek and nape. The dorsal profile of the head and snout in L. vellicans
apparently differs from that of P. victoriae, in being straighter and more steeply
sloping.
Amongst the Haplochromis species of Lake Victoria, P. victoriae, has some structural
affinity with Haplochromis chilotes. Both species have short and narrow jaws,
thickened lips and a certain similarity of dentition. For example, the teeth of H.
chilotes are slender and elongate anteriorly, few in number and arranged in an acute
dental arch. However, the disproportionately long and procurrent anterior teeth of
P. victoriae, and the hypertrophied lips of H. chilotes, immediately serve as diagnostic
characters. The phylogeny of H. chilotes, is at present, obscure, but the species
would seem more closely related to the H. crassilabris species group in Lake Victoria,
than to Paralabidochromis.
DISCUSSION
It is clear from Regan's analysis of the Lake Victoria Cichlidae, that he did not
consider the monotypic genera far removed phylogenetically from certain Haplo-
chromis species, a view which is strengthened by the additional data now available.
Although the monotypic genera are readily defined by trenchant characters they
retain fundamental affinities with the Haplochromis species of Lake Victoria. But
the morphological differentiation which these genera have undergone creates an
impression of greater divergence than is shown by other adaptive groups within the
Lake Victoria species flock. Analysis of the diagnostic characters of the genera
described here, shows that, in any one genus, the anatomical characters of the head
are functionally related to the dentition. Further, the ontogenetic basis for most of
these characters is probably attributable to differential growth and not to any
large-scale qualitative change. Thus the first evolutionary steps involved may well
have been relatively simple and similar, in the earliest stages, to those which initiated
the often slight differences characterizing the greater number of Haplochromis species.
330 MONOTYPIC GENERA OF CICHLID FISHES IN LAKE VICTORIA
Despite fundamental inter-relationship, three of the four monotypic genera
exhibit considerable inter-generic divergence in the characters affected by the peculiar
development of their dentition. The degree of divergence and the means by which
it has been effected in Macropleurodus, Platytaeniodus and Hoplotilapia are such
that the genera cannot be related inter se. That is to say, these genera represent
three independent evolutionary offshoots from the basic Haplochromis stock. The
fourth genus, Paralabidochromis, shows least departure from the generalized Haplo
chromis condition, but must also be looked upon as having evolved independently.
A broad outline of the probable phyletic relationships within the Haplochromis
species flock is perhaps necessary before attempting further discussion of the
monotypic genera.
There are some Lake Victoria Haplochromis species which, when seen in isolation
or when known only from a few specimens, might seem almost as distinctive as the
monotypic genera. It is possible, however, to relate these outstanding species to
others more typical of their particular adaptive group (mollusc eaters, piscivores,
epiphytic-algal grazers, etc.). Furthermore, when larger series of specimens are
examined, infra-specific variation is sufficiently high to reduce considerably any
apparent inter-specific gap.
With few exceptions, specific differences amongst the Haplochromis are quantitative
and generally consequent upon the differential growth of various syncranial parts.
Qualitative differences, on the other hand, are usually those which can be related to,
and are used in defining, adaptational groups. At both inter- and intra-group levels,
qualitative and quantitative differences tend to be small and intergrading.
It is in both qualitative and quantitative characters that the monotypic genera
depart most markedly from Haplochromis but, unlike the inter-group differences
within the Haplochromis flock, the morphological gap is clear cut, and remains so
even when large series of specimens are examined.
On this interpretation, Hoplotilapia and Platytaeniodus probably examplify the
phenomenon of " quantum evolution " as described by Simpson (1944 and 1953). It
is difficult to suggest any ancestry for these genera nearer than a present-day species
group, viz. the H. crassilabris complex. Even at this level relationship is extremely
tenuous, and based only on similarity of skull form, in itself probably an adaptational
character and therefore of doubtful phylogenetic significance. The jaws and dental
pattern are so dissimilar in Hoplotilapia and Platytaeniodus that one must consider
the genera separately and not as elements of a single lineage.
Macropleurodus is less readily regarded as being a product of quantum evolution,
and will be discussed later.
Simpson (1944) has noted that quantum evolution is usually associated with a
shift from one adaptive zone to another, and that the interzonal populations or
species would be relatively ill-adapted, unstable and short-lived. Thus, morpho-
logical discontinuity is generally observed between the parental and divergent
lineages.
The morphological discontinuity existing between Platytaeniodus and Hoplotilapia,
and between these two genera and Haplochromis has been demonstrated ; it is the
more regrettable then that pertinent ecological data for P. degeni are both inadequate
MONOTYPIC GENERA OF CICHLID FISHES IN LAKE VICTORIA 331
and confusing, so that it is difficult to equate the apparent morphological specializa-
tion of this species with any particular ecological niche. The food of P. degeni is
virtually unknown, although inorganic material and scant but varied organic
remains suggest bottom-feeding habits. A multi-serial and concentrated dentition,
such as that of P. degeni may possibly have adaptive value, particularly if food must
be freed from the substrate or if it requires trituration before digestion can be effected.
(It should be noted that no Haplochromis has consistently yielded such baffling
residua in the gut.)
Hoplotilapia retrodens is somewhat better understood. In this species food is
predominantly Molluscan. The functional significance of the jaw structure and
dental pattern in Hoplotilapia, particularly with regard to its observed diet of thin-
shelled bivalves, has been discussed above (p. 325). From these characters and the
predominance of Mollusca over the other food organisms it would seem that the
essential elements of quantum evolution are fairly well defined in H. retrodens. That
is to say, the species shows both morphological discontinuity and entry into an
adaptive niche different from that of the presumed parental stock.
There is, of course, the possibility that Platytaeniodus and Hoplotilapia represent
an early stage in quantum evolution and that these species may be relatively ill-
adapted to existing ecological conditions. Their further evolutionary development
or, alternatively, extinction, will therefore depend upon environment change effecting,
or failing to effect, the realization of characters which at present could only be
considered prospective adaptations (sensu Simpson, 1953, p. 188).
The slight and often indeterminable adaptive differences between Haplochromis
species occupying the same ecological zone, together with numerous instances of
inter-specific overlap in feeding habits, would seem to suggest that there is, and has
been, only slight selection-pressure acting through competition for food. Thus the
continued existence of Platytaeniodus and Hoplotilapia, like that of many closely
related Haplochromis species, could be attributed to a period of decreased selection
pressure.
Although Macropleurodus, like Platytaeniodus and Hoplotilapia, is separated from
Haplochromis by a clearly defined morphological gap, the gap is of lesser degree.
Whereas P. degeni and H. retrodens exhibit unique jaw morphology and dental
patterns without departure from Haplochromis in tooth form, the basic dental
pattern of M. bicolor is foreshadowed in two Haplochromis species, as is the form of
the dentary. Only in tooth form and shape of the premaxilla does M. bicolor show
great differentiation from H. prodromus. The latter species is morphologically
closely related to H. sauvagei, a smaller species, which in turn shows departure from
the basic Haplochromis type towards H. prodromus. In H. sauvagei and H. prodromus
the outer teeth are relatively stout and, anteriorly, there are several rows of inner
teeth ; the neurocranium and dentary of these species also approach the M. bicolor
condition. Both species feed on Mollusca and Insect a, the very slight differences in
the feeding habits of H. prodromus and H . sauvagei being attributable to the smaller
size of H. sauvagei.
It is tempting, therefore, to consider the members of the series H. sauvagei-
H. prodromus-M. bicolor as representing stages of a lineage, although the possibility
332 MONOTYPIC GENERA OF CICHLID FISHES IN LAKE VICTORIA
of independent evolution of the same adaptive characters, cannot be ignored. Until
more critical evidence is available for the phyletic relationship of these species, they
may be regarded either as separate end points of different lineages, or as a " stufen-
reine", or single phyletic line. The three species do illustrate the gross anatomical
and functional stages through which Macropleurodus could have evolved, although
the transition in shape of teeth from a typical Haplochromis to Macropleurodus is
not represented in any living species.
The isolated position of Paralabidochromis in relation to the Haplochromis species
of Lake Victoria was commented upon above.
Little information is available on the evolutionary relationships of Paralabido-
chromis. Although taxonomically isolated from Haplochromis the genus is nearer
the Haplochromis stem than either Hoplotilapia, Platytaeniodus or Macropleurodus.
In Lake Nyasa, Trewavas (1935 and 1949) considers Labidochromis vellicans as
belonging to a group of nine genera (which group excludes Haplochromis), that,
although lacking an absolute character to distinguish them, are more closely related
to each other than to any other genus. Such grouping is impossible for the monotypic
genera of Lake Victoria. Within this lake the divergent genera of the Haplochromis
group must be considered as being distinct from one another, as well as from the
parental stock.
SUMMARY
(1) The genera Macropleurodus Regan, 1922, Platytaeniodus Boulenger, 1906, and
Hoplotilapia Hilgendorf, 1888, are redefined on the basis of new and fairly extensive
collections. Similarly, the species M. bicolor (Blgr.) 1906, P. degeni Boulenger, 1906,
and H. retrodens Hilgendorf, 1888, are redescribed.
(2) Generic and specific characters are discussed, with particular regard to onto-
genetic changes. Information gained from small specimens shows that three species
previously considered as Haplochromis must be added to the synonymies of M.
bicolor and P. degeni.
(3) Comparative anatomical and osteological studies of the head indicate that
Hoplotilapia and Platytaeniodus are not closely related to any extant Haplochromis
species in Lake Victoria. Morphological stages leading to the syncranial type found
in Macropleurodus are, however, represented by two endemic Haplochromis species.
On the basis of syncranial morphology, it is clear that the monotypic genera are not
closely related inter se.
(4) Apparently sex-limited polychromatism occurring in Macropleurodus and
Hoplotilapia is described and discussed.
(5) A fifth monotypic genus, Paralabidochromis victoriae is described. This genus
exhibits remarkably close morphological parallelism with Labidochromis vellicans
from Lake Nyasa.
(6) Locality lists and notes on the ecology of the genera are given, together with
observations on the feeding habits of Macropleurodus and Hoplotilapia.
(7) The evolutionary relationships of the four genera are discussed. It is suggested
that Hoplotilapia and Platytaeniodus may represent the products of low-level
quantum evolution.
MO NOT Y PIC GENERA OF CICHLID FISHES IX LAKE VICTORIA 333
A C K N O W L E D G M E N T S
I wish to acknowledge my gratitude and thanks to the Trustees of the British
Museum (Natural History) for facilities afforded me during the tenure of a Colonial
Fisheries Research Studentship ; to Professor L. Bertin of the Museum National
d'Histoire naturelle, Paris, for allowing me to study type specimens of Lake Victoria
Cichlidae described by Pellegrin ; to Dr. Delfa Guiglia of the Museo Civico di Storia
Naturale, Genoa, for the courtesies mentioned in the text ; to Dr. Kurt Deckert,
Zoologisches Museum, Berlin, for attempting to locate the type specimen of Hoplo-
tilapia retrodens ; to the officers of the Lake Victoria Fisheries Service who collected
much valuable material ; to my colleagues Miss R. H. Lowe and Dr. P. S. Corbet
for many discussions and to the latter for his invaluable identifications of material
from gut-contents ; and to Mr. Denys W. Tucker, for his very helpful criticism of
the manuscript. I am especially indebted to Dr. Ethelwynn Trewavas for the
interest she has shown in my work, and for much helpful information and advice.
REFERENCES
This list includes only those references which occur in the general sections of the text. All
others appear under the corresponding species headings:
Cott, H. B. 1952. In East African Fisheries Research Organization Annual Report. East
Africa High Commission, Nairobi (p. 21).
Graham, M. 1929. A report on the Fishing Survey of Lake Victoria, 1927-1928, and appendices.
Crown Agents, London.
Greenwood, P. H. 1954. On two cichlid fishes from the Malagarazi river (Tanganyika),
with notes on the pharyngeal apophysis in species of the Haplochrotnis group. Ann. Mag.
nat. Hist. (12) 7 : 401.
Regan, C. T. 1920. The classification of the fishes of the family Cichlidae. I. The Tangan-
yika genera. Ann. Mag. nat. Hist. (9) 5 : 33.
■ — — 192 1. The cichlid fishes of Lake Nyasa. Proc. zool. Soc. Lond. 675.
— ■ — 1922a. The cichlid fishes of Lake Victoria. Proc. zool. Soc. Lond. 157.
1922&. The classification of the fishes of the family Cichlidae. II. On African and
Syrian genera not restricted to the Great Lakes. Ann. Mag. nat. Hist. (9) 10 : 249.
Simpson, G. G. 1944. Tempo and Mode in Evolution. New York.
1953- The Major Features of Evolution. New York.
Trewavas, E. 1933. Scientific results of the Cambridge expedition to the East African lakes,
1930-1. II. The cichlid fishes. /. linn. Soc. {Zool.) 38 : 309.
T 935- A synopsis of the cichlid fishes of Lake Nyasa. Ann. Mag. nat. Hist. (10) 16 : 65.
— — 1949. The origin and evolution of the cichlid fishes of the great African lakes, with
special reference to Lake Nyasa. Rapp. 13 e Congres Intern. Zool., Section 5B, 365. Paris.
PRINTED IN GREAT BRITAIN BY
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- Z MAH mb
A REVISION OF THE HYDROID
GENUS PERIGONIMUS
M. SARS, 1846
ON THREE NORTHERN SPECIES
OF HYDRACTINIA
WILLIAM J. REES
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 3 No. 8
LONDON: 1956
A REVISION OF THE HYDROID GENUS
PERIGONIMUS M. SARS, 1846
ON THREE NORTHERN SPECIES OF
HYDRACTINIA
BY
WILLIAM J. REES
Pp. 335-362 ; Pis. 11-12 ; 2 Text-figures
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 3 No. 8
LONDON: 1956
THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, is
issued in five series, corresponding to the Departments
of the Museum, and an Historical Series.
Parts will appear at irregular intervals as they
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pleted within one calendar year.
This paper is Vol. 3, No. 8 of the Zoological Series.
PRINTED BY ORDER OF THE TRUSTEES OF
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Issued February, 1956 Price Ten Shillings.
A REVISION OF
THE HYDROID GENUS PERIGONIMUS
M. SARS, 1846
By WILLIAM J. REES
SYNOPSIS
During the past century nearly forty hydroids have been referred at one time or another
to the genus Perigonimus M. Sars. The type species was demonstrated to be a Bougainvillia
in 1938, and the status of all the other " Perigonimus " spp. is now considered and their position
determined in accord with modern ideas on classification. They are distributed between the
Orders Anthomedusae (Athecata), Leptomedusae (Thecata) and Limnomedusae, the majority
being referred to four families of the first-mentioned Order.
Some material of Perigonimus vestitus, determined by Allman, has been re-discovered in the
British Museum and is found to belong to Leuckartiara octona (Fleming) . The original drawings
of Joshua Alder's Perigonimus linearis, also in the Museum, reveal the presence of oral tentacles
on the manubrium of the newly-liberated medusa, thus confirming that Bougainvillia flavida
Hartlaub is identical with it.
INTRODUCTION
In 1938, I published a partial revision of Perigonimus in so far as it affected the
groundwork for a monograph of British medusae, later published by F. S. Russell
( I 953)- The earlier revision facilitated the establishment of common names for
both the hydroid and the medusa of British species, but did not indicate the position
of many extra-territorial ones whose status needed to be clarified. In this revision
the position of these remaining species is considered and the conclusions reached
earlier for others are re-stated briefly.
Michael Sars (1846) described a new handsome hydroid from the Norwegian
coast under the name Perigonimus muscoides and also gave a brief, but inadequate,
description of the medusa which is liberated with four tentacles. Later Allman
(1864a), in his revision of Bougainvillid and other hydroids, referred a number of
them which could not be placed in Bougainvillia (because they did not produce
Bougainvillid medusae) to this genus of Sars on the assumption that they were
cogeneric. Thus Perigonimus came to represent a group of Bougainvillia-like
hydroids, which, either did not give rise to Bougainvillia medusae or had fixed
gonophores. Gradually the picture emerged that the medusae of most of these
non-Bougainvillia species belonged to another family, the Pandeidae. Hartlaub
(1897 and 1914) demonstrated that Perigonimus repens was the hydroid of " Tiara
pileata", now known as Leuckartiara octona (Fleming), and that another Pandeid,
Halitholus cirratus Hartlaub, also had a Perigonimus hydroid. More recently Rees
& Russell (1937) were able to link the Perigonimus serpens kinds of hydroids with
two species of medusae also of this family. Further revision of Perigonimus was
zool. 3, 8,
338 A REVISION OF THE HYDROID GENUS PERIGONIMUS M. SARS
made possible by the discovery that the type species of Perigonimus gave rise to a
Bougainvillia medusa (Rees, 1938). Consequently Perigonimus became a synonym
of Bougainvillia and no longer available as a name for hydroids which could not be
placed in the same genus or even the same family.
As Amphinema (i.e., Stomotoca) had already been established by Rees & Russell
(1937) for the hydroids as well as the medusae of this genus, Leuckartiara (already
re-established for the medusa phase) was adopted for the appropriate hydroids, and
Rhizorhagium M. Sars re-established for non-medusa bearing species.
As already mentioned this left a number of " Perigonimus " species either not
satisfactorily classified or left out of consideration, and, in the brief notes below,
it has been thought desirable to indicate their present status and where possible to
add new information.
THE SPECIES WHICH HAVE BEEN REFERRED TO PERIGONIMUS.
Perigonimus abyssi G. O. Sars
Perigonimus abyssi G.O. Sars, 1873, Forh.Vidensk. Selsk. Krist., 1873 : 126, Tab. v, figs. 27-30.
The newly liberated medusa of this minute species becomes free with two opposite
tentacles and two partly developed tentacles on the other perradial bulbs (Rees,
1938). It is a Pandeid and has been referred provisionally to Leuckartiara Hartlaub
pending further knowledge of its possible relationship with medusae of the genera
Leuckartiara, Neoturris or Catablema.
Perigonimus antarcticus Hickson & Gravely
Perigonimus antarcticus Hickson & Gravely, 1907, Nat. Antarct. Exped., 1901-1904, Nat. Hist.,
3 : 1, pi. i, figs. 1-3 ; pi. iv, fig. 32.
This species was placed in Atractylis (synonymous with Bougainvillia) by Van-
hoffen (1910) and in a new genus Gravely a by Totton (1930), but its affinities lie
with Rhizorhagium M. Sars, to which genus it was referred by Stechow (1919) and
by Rees (1938). This last mentioned genus seems to be well constituted for simple
Bougainvillid hydroids with fixed gonophores, but its relationship to Aselomaris
Berrill and Garveia Wright will have to be considered in a wider revision of the
Bougainvillidae.
Perigonimus apicatus Brooks
Perigonimus (Stomotoca) apicata Brooks, 1884, Zool. Anz., 7 : 711.
This is the name given by Brooks to the hydroid of Stomotoca apicata, now known
as Stomotoca dinema (Peron & Lesueur). Mayer (1910, p. 113) referred this hydroid
to 5. rugosa Mayer, but the absence of an apical projection in the newly liberated
medusa points to 5. dinema, as does the name used by Brooks for his hydroid.
Perigonimus bitentaculatus (Wright)
Atractylis bitentaculata Wright, 1867, Proc. R. Phys. Soc. Edinb., 3 : 45, pi. i, fig. 5.
This species was referred provisionally to Perigonimus by Hincks (1868), but
Wright himself gives the best clue as to the relationships of his species in the following
A REVISION OF THE HYDROID GENUS PERIGONIMUS M. SARS 339
sentence : " They have the habit, like that of Lar (Gosse), of quickly and repeatedly
bending down the body until the mouth is brought close to the surface on which
the zoophyte grows."
This species has not been reported since it was found by Wright in a Pecten shell
dredged from the Firth of Forth near Inch Keith. The characters of the hydranth
with its two opposite tentacles strongly suggest that the hydroid is related to Pro-
boscidactyla (Lar of Gosse) and should be placed in the same family. The possibility
should not be overlooked that this species may prove to belong to Pochella polynema
Hartlaub, the only other medusa of this family known to occur in British waters.
I do not propose to create a new genus for this species, but merely note that its
position appears to be in the family Proboscidactylidae of the Order Limnomedusae.
Perigonimus cidaritis Weismann
Perigonimus cidaritis Weismann, 1883, Entstehung der Sexualzellen bei den Hydromedusen, Jena,
p. 117, pi. xii, fig. 10— 11.
This hydroid has a well-marked pseudohydrotheca in which the bases of the
tentacles are protected in tube-like extensions of the perisarc as in Bimeria and
Thamnostoma. As the medusa is a Bougainvillid closely related to Thamnostoma
russelli Rees and there is very little difference in the hydroids, P. cidaritis has been
referred to Thamnostoma by Rees (1938).
Perigonimus cirratus Hartlaub
Perigonimus cirratus Hartlaub, 1914, Nord. Plankt., Lief. 17, XII : 274, fig. 234.
Hartlaub (1914) gave the name P. cirratus to the hydroid of the medusa Halitholits
cirratus which he described in the same memoir, presumably for the reason that it
was then customary to adhere to a dual system of classification. There are now no
legitimate reasons why the hydroid should not also take the name Halitholus.
Perigonimus coccineus (Wright)
Atractylis coccinea Wright, 1861, Ann. Mag. Nat. Hist., (3) 8 : 130.
E. S. Russell (1907) is the only author who has reported this species since it was
originally described by Wright. In 1938 I suggested that the species might prove to
be identical with Rhizorhagium roseum.
Perigonimus confertus (Alder)
Eudendrium ? confertum Alder, 1856, Trans. Tyneside Nat. Fid. CI., 3 : 103, pi. iii, figs. 5-S.
Allman (1859) placed this species in Dicoryne because of the peculiarities of the
gonophores, an assessment which has been generally approved. Recently Vervoort
(1946) used the name Perigonimus confertus for the first time for this species, although
presumably aware that this name is synonymous with Bougainvillia and therefore
no longer available. The name Dicoryne conferta has been the established name of this
species since 1859 and I see no reason to change it.
340 A REVISION OF THE HYDROID GENUS PERIGONIMUS M. SARS
Perigonimus decorans Schneider
Perigonimus decorans Schneider, 1897, Zool. Jb. Syst., 10 : 479.
Schneider's description is too inadequate even to indicate in what family this
hydroid could be placed.
Perigonimus formosus Fewkes
Perigonimus formosus Fewkes, 1889, Bull. Essex Inst., 21 : 102, no figure.
Some small thumb-nail sketches of this species have been given by Fraser (1937,
p. 35, pi. v, fig. 20) ; they indicate that P. formosus should be placed in Rhizorhagium
M. Sars. A new description, especially of the gonophores, is much needed to determine
the relationship of the species to R. roseum.
Perigonimus gelatinosus Duerden
Perigonimus gelatinosus Duerden, 1895, Sci. Proc. R. Dublin Soc, n.s., 8 : 325, 327, pi. xiv,
figs. 2-3.
Duerden's species is not identifiable with certainty with any known British species,
although there is a possibility that it is identical with Leuckartiara octona as supposed
by Hartlaub (1914). The complete retraction of the hydranth within the pseudo-
hydrotheca could be due to poor preservation and by itself is no indication of specific
differences.
Perigonimus georginae Hadzi
Perigonimus georginae Hadzi, 1913, Rad. Jug. Akad. Znan. Umj., 200 : 98, figs. 7, 10 and 11.
The original description by Hadzi is not very satisfactory. He appears to have
confused at least two species. His figure 7 is a Stomatoca hydroid, his figure 8 repre-
sents a Bougainvillia, and the newly liberated medusa, although combining some of
the features of two species of Stomotoca medusae, cannot be referred with certainty
to either 5. rugosa Mayer or to 5. dinema (Peron & Lesueur). Until the Adriatic
medusae have been re-investigated it seems preferable to let this species retain its
identity as Stomotoca georginae. The name Dinema applied by Hadzi to the medusa
is a synonym of Leuckartiara, a member of a different sub-family.
Perigonimus (?) inflatus Duerden
Perigonimus (?) inflatus Duerden, 1895, 5a. Proc. R. Dublin Soc, n.s., 8 : 329, pi. xiv, fig. 4.
This little known species does not seem to have been reported since Duerden's
original description. It is here referred provisionally to Rhizorhagium M. Sars.
Perigonimus jonesii Osborn & Hargitt
Perigonimus jonesii Osborn & Hargitt, 1894, Amer. Nat., 28 : 27, figs. 1-12.
I have followed Hartlaub (1914) in placing this species in the synonymy of
Leuckartiara octona (Fleming). The colony described by Osborn & Hargitt compares
favourably with those found on Scaphander lignarius in British waters.
A REVISION OF THE HYDROID GENUS PERIGONIMUS M. SARS 341
Perigonimus linearis (Alder)
Atractylis linearis Alder, 1862, Trans. Tyneside Nat. Fid. CI., 5 : 230, pi. x, figs. 1-3.
Alder's Atractylis linearis was tentatively referred to Perigonimus byAllman (1864a).
The newly liberated medusa has a very thick jelly and a small subumbrellar cavity,
and in this it resembles Bougainvillia flavicla Hartlaub (1897), an opinion later
expressed by Hartlaub himself (1911). The published drawings of Alder do not
suggest any oral tentacles on the manubrium of the medusa, but fortunately his
original drawings (from which the engraving for his paper (1862, pi. x, figs. 2 and 3)
was prepared) are preserved in the British Museum (Nat. Hist.), having been acquired
by Canon A. M. Norman and later bequeathed by him. These drawings do show the
oral tentacles faintly and so the two species can be regarded as identical.
Russell (1953) does not venture an opinion on the status of P. linearis or its medusa,
but suggests that B.flavida may prove to be the hydroid of Bougainvillia britannica.
Until this relationship can be demonstrated I propose to retain Alder's species under
the name Bougainvillia linearis (Alder, 1862).
Perigonimus maclovianus (Lesson)
Perigonimus maclovianus Vanhoffen, 1910, Dtsch. Sudpolar-Exped., 11 : 272, fig. loa-d (hydroid).
The medusa of this species was described under the name Cyanaea bougainvillii
by Lesson (1830), who later made it the type species of Bougainvillia under the name
B. macloviana (1836). This is the name in common use to-day for both hydroid and
medusa, although Vanhoffen (1910), who first described the hydroid, referred it to
Perigonimus. Both Stechow (1919, p. 21) and Kramp (1928, p. 50) pointed out that
Bougainvillia was the correct generic name for both hydroid and medusa.
Perigonimus minutus Allman
Perigonimus minutus Allman, 1863, Ann. Mag. Nat. Hist., (3), 11 : 11.
Allman's Perigonimus minutus has been referred to Leuckartiara oclona both by
Hartlaub (1914) and Rees (1938). Allman's description and figures (1871-2, pi. xi,
figs. 4-6) correspond closely with the slender polyps such as are commonly found on
Scaphander lignarius (Rees, 1938, p. 14 fig. 3d-f).
Perigonimus miniatus (Wright)
Atractylis miniata Wright, 1863, Proc. R. Phys. Soc. Edinb., 2 : 351.
Wright's Atractylis miniata was referred provisionally to Perigonimus by Hincks
(1868). The reproduction of the species is not known, but the description suggests that
Wright was describing a colony of Rhizorhagium and R. roseum in particular.
Perigonimus multicornis Allman
Perigonimus multicornis Allman, 1874, Nature, Lond., 11 : 179.
A re-examination of Allman's type material in the Zoological Museum, Copenhagen
by Kramp (1926) revealed that the species was Eudendrium ramosum.
342
A REVISION OF THE HYDROID GENUS PERIGONIMUS M. SARS
Perigonimus muscoides M. Sars
Perigonimus muscoides M. Sars, 1846, Fauna Littoralis Norvegiae, Heft. 1 : 8, Tab. 1, figs. 19-21.
This is the type species of Perigonimus and the elucidation of its life history by Rees
(1938, p. 2) revealed that its medusa was a Bougainvillia, B. nordgaardi (Browne,
1903). Perigonimus M. Sars, 1846, thus falls into the synonymy of Bougainvillia
Lesson 1836 and the name Bougainvillia muscoides becomes applicable to both
hydroid and medusa as demonstrated in the above paper.
Perigonimus muscus Allman
Perigonimus muscus Allman, 1863 ; Ann. Mag. Nat. Hist. (3), 11 : 12.
This species was referred to Bougainvillia by Allman (18640). It was considered to
be synonymous with Bougainvillia ramosa by Hartlaub (1911) and this view was
confirmed by the studies of Brink (1925) who demonstrated that colonies passed
through a muscus stage when young before reaching the stage typified by the usual
ramosa kind of colony.
Perigonimus (?) nanellus Stechow
Perigonimus (?) nanellus Stechow, 1919, Zool. Jb. Syst., 42 : 14, fig. C.
I consider this species to be a Bougainvillia, possibly representing a well formed
colony of one of the common species, and pending further knowledge of its life
history it is now referred to this genus as B. nanellus.
Perigonimus napolitanus Hargitt
Perigonimus napolitanus Hargitt, 1904, Mitt. zool. Sta. Neapel, 16 : 571, taf. 22, fig. 25.
This species is not recognizable.
Perigonimus nudus Stechow
Perigonimus nudus Stechow, 1919, Zool. Jb. Syst., 42 : 16, fig. D.
The form of the hydranth as noted by Stechow resembles those of the medusa-
bearing Campanopsinae of the family Haleciidae and the relationships of the species
undoubtedly lie with this group rather than with any Athecate (Anthomedusan)
group.
Perigonimus ? nutans Hincks
Perigonimus (?) nutans Hincks, 1877, Ann. Mag. Nat. Hist., (4), 19 : 149, pi. xii, fig. 1.
Rees & Russell (1937, p. 71) regard this species as possibly a young polyp of the
Perigonimus serpens type ; this kind of hydroid belongs to the genus Amphinema.
There are two species in British waters, A. dinema and A. rugosum, and these are so
alike that it is not possible to assign this species to one or the other.
Perigonimus palliatus (Wright)
Atractylis palliata, Wright, 1861, Ann. Mag. Nat. Hist. (3), 8 : 129, pi. iv, figs. 6 and 7.
In this species the pseudohydrotheca is large and dilated, but I do not think it has
any special significance as a diagnostic character. The medusa, according to Wright
A REVISION OF THE HYDROID GENUS PERIGONIMUS M. SARS 343
(1861) " resemble exactly those of Atractylis (Perigonimus) repens ", that is, Leuckarti-
ara octona (Fleming). Wright's colony was on a shell inhabited by a hermit crab
Eupagurus bernhardus and the size of the pseudohydrotheca can be attributed to
habitat and feeding conditions as in the other colonies of Leuckartiara octona living
under similar conditions (see Rees, 1938, p. 16, fig. 5). I regard the two species as
identical.
Perigonimus pugetensis Heath
Perigonimus pugetensis Heath, 19 10, Biol. Bull. Woods Hole, 19 : 74, figs. 1 and 2.
This small hydroid was found growing on a fish, Hypsagonus quadricornis (C. & V.) ;
it was found that its newly liberated medusa was a Pandeid like Leuckartiara octona.
Hartlaub (1914) doubted whether the species was distinct from L. octona, and
the latter has since proved to be a cosmopolitan species. Rees (1938) found a colony
of L. octona growing on a fish, Agonus cataphr actus, and this colony compared very
favourably with that of Heath. It is now proposed to regard P. pugetensis as a syno-
nym of L. octona.
Perigonimus pusillus (Wright)
Eudendrium pusillum Wright, 1857, Edinb. New Phil. J., N.s., 6 : 84, pi. ii, fig. 8, 9.
Stechow (1919) revived the name pusillus for the better known name Perigonimus
repens (Wright, 1858). It appears that Wright changed the name of his species to
repens on the receipt of a paper by M. Sars (1857) in which he described a species of
Halecium under the name Eudendrium pusillum.
Wright's Species falls into the synonymy of Leuckartiara octona (Fleming).
Perigonimus quadritentaculatus (Wright)
Atractylis quadritentaculata Wright, 1867, Proc. R. Phys. Soc. Edinb. 3 : 45, pi. i, fig. 6.
This creeping form with long stolons and sessile, four tentacled hydranth is possibly
a juvenile Bougainvillid, which cannot be identified with certainty with any particular
species.
f Perigonimus robustus Fraser
? Perigonimus robustus Fraser, 1938, Rep. Hancock Pacific Exped., 4, No. 1 ; 17, pi. iii, fig. 12.
In the absence of information on the gonophore this species cannot be satisfactorily
classified, and is therefore provisionally placed in Bougainvillia.
Perigonimus roseus (M. Sars)
Rhizorhagium roseum M. Sars, 1877, Fauna Littoralis Norvegiae, Heft 3 : 28, Tab. ii, figs. 37-43.
The genus Rhizorhagium was created for this species by Sars, and although the
type species was referred for a time to Perigonimus by Bonne vie (1899) and others,
the genus was re-instated by Rees (1938).
Perigonimus sarsii Bonnevie
Perigonimus sarsii Bonnevie, 1901, Bergens Mus. Meeresfauna von Bergen, Heft I, p. 6.
This species has been referred to Rhizorhagium by Rees (1938) and there is a
distinct possibility that the species is founded on a colony of R. roseum, growing over
the stems of another hydroid,
ZOOL. 3, 8,
344 A REVISION OF THE HYDROID GENUS PERIGONIMUS M. SARS
Perigonimus schneideri Motz-Kossowska
Perigonimus schneideri Motz-Kossowska, 1905, Arch, Zool. exp. gen., (4), 3 : 72, fig. VI.
Stechow (1922) created a new genus Perarella for this species and indicated that
its position was not in the Hydractinidae but rather in the Bougainvillinae near
Atractylis.
Since then Komai (1931) has described the hydroid of the medusa Cytaeis japonica,
to which the present species is clearly related. Both species have the following
features in common : a reticulate anastomozing network of stolons, a short collar
round the base of the hydranth, the hydranth itself long and tubular, of the Podo-
coryne type, and the gonophores borne singly on the hydrorhiza.
I propose to retain Perarella for Cytaeid hydroids in which there are fixed gonophores
and to place the genus in the family Cytaeidae, L. Agassiz, 1862. Agassiz's definition
excludes unrelated forms such as Podocoryne (including Lymnorea), Turritopsis and
Oceania, which Uchida (1927) and others have placed in the family.
While possessing distinct oral tentacles on the manubrium of the medusa as in the
Bougainvillidae, the presence of simple perradial tentacles together with the characters
of the hydroid justify the existence of a separate family for Cytaeis and Perarella — a
family in which the combined characters of hydroid and medusa present some
characters in common with both the Hydractinidae and the Bougainvillidae, but
none of their specialized features.
Perigonimus serpens Allman
Perigonimus serpens Allman, 1863, Ann. Mag. Nat. Hist. (3), 11 : 10.
Rees and Russell (1937) have referred this species to Amphinema dinema following
the elucidation of the life history of this medusaand its hydroid. Allman (1871-2) clear-
ly indicated in his illustrations of the medusa that he was dealing with this species
and not with the closely related A . rugosum.
Perigonimus sessilis (Wright)
Eudendrium sessile Wright, 1857, Edinb. New Phil. J, n.s., 6 : 90, pi. iii, figs. 16-17.
This is the sessile form of Leuckartiara octona (Fleming). When the colony is
growing on an uneven, constantly abraded surface the hydranths remain sessile in
protected hollows or grooves and do not progress beyond the sessilis stage (Rees,
1938).
Perigonimus steinachi Jickeli
Perigonimus steinachi Jickeli, 1883, Gegenbaurs Jb., 8 : 617, pi 27, figs. 1-9.
Jickeli gives a totally inadequate description of the morphology of his hydroid
(although he discusses its histology at length) and it is not possible to indicate what
it might be.
Perigonimus sulfureus Chun
Perigonimus sulfureus Chun, 1889, 5. B. Akad. Wiss. Berlin, 1889 (2) : 524.
Stechow (1921 and 1923) created a new genus Perigonella for this species and placed
it with the Hydractinians near Stylactella. Steche (1906) figures a well-developed
A REVISION OF THE HYDROID GENUS PERIGONIMUS M. SARS 345
medusa bud with conical bulbs with tentacles and manubrium devoid of oral tentacles,
but we have insufficient information to assign this species to a family.
Perigonimus vagans Thornely
Perigonimus vagans Thornley, 1908, /. Linn. Soc. Zool., 31 : 81, pi. ix, fig. I.
This species is now referred provisionally to Bougainvillia pending the re-discovery
of the hydroid. Thornely's description is insufficient to permit any other suggestions.
Perigonimus vestitus Allman
Perigonimus vestitus Allman, 18646, Ann. Mag. Nat. Hist., Ser. (3), 14 : 57.
In 1846 Allman described a new Perigonimus, P. vestitus from the Firth of Forth,
in which, to quote his 1872 diagnosis the " posterior part of the body [of the hydranth],
invested by the rough perisarc, which is thence continued as a delicate, smooth
membrane over the remainder of the body nearly as far as the mouth ". His figures
(1871, pi. xi, figs. 1-3) show the pseudohydrotheca (that is the perisarcal investment
of the hydranth) to extend anteriorly beyond the tentacles close to the mouth. There
is no suggestion that this perisarc is perforated to allow the tentacles to project (as
they do) and no suggestion of any tubular perisarcal sheathing encircling the base of
any tentacle. This seems to be one of the few occasions on which this far-sighted
pioneer of hydroid systematics failed to make a satisfactory description.
Stechow (1919) referred P. vestitus to the medusa genus Cytaeis presumably on an
assumed relationship between the hydroid and medusa of vestitus with Perigonimus
cidaritis. There were no real grounds for this, as the figures demonstrate that P.
vestitus gives rise to a typical Pandeid medusa, so that it must be stated that the
species is related to genera like Leuckartiara, Amphinema and Halitholus.
In Perigonimus vestitus too, the bases of the tentacles are not enclosed in perisarcal
tubes extending from the pseudohydrotheca itself. Such a feature is characteristic
of Bimeria, Thamnitis and Thamnostoma, and the medusa of the latter clearly has
affinities with the Bougainvillidae rather than the Pandeidae. Cytaeis, which Stechow
erroneously associated with these Bimerid hydroids, belongs to none of these families.
Hartlaub (1914) in his revision of the Tiaridae had listed Perigonimus vestitus
under the synonymy of Leuckartiara octona, but I was reluctant to follow him in
my study of variation in the hydroid of this species (1938) in view of the diagnosis
given by Allman. Fortunately, material of Perigonimus vestitus determined by Allman,
growing on Lepidochiton cinereus, has recently been found in the collections of the
British Museum (B.M. No. 1877.4. 12.31).
A re-examination reveals that although the soft parts have disappeared the
chitonous perisarc of the stems and pseudohydrothecae are still present. The latter
are very similar in appearance to those that I described for a colony of Leuckartiara
octona (Fleming) on Corystes cassivelaunus (Rees, 1938, p. 16, fig. 5). There is no
suggestion of any perisarcal sheaths such as are found in Bimeria. The colonies
exhibit no features that could not be found in a well-developed colony of Leuckartiara
octona, and the characters of the medusa, as described by Allman, support this. For
346 A REVISION OF THE HYDROID GENUS PERIGONIMUS M. SARS
these reasons the species vestitus is placed in the synonymy of Leuckartiara octona
(Fleming).
Perigonimus yoldiae-arcticae Birula
Perigonimus yoldiae-arcticae Birula, 1897, Annu. Mus. Zool. Acad. St. Petersb. 2 : 86, pi. x, fig. 3.
Jaderholm (1909) gives an excellent figure of this hydroid. I have examined
material from the following localities :
Zoologiske Museum, Copenhagen.
Kandalakscha, on Yoldia arctica, 24 fm. (from St. Petersbourg Museum).
Riksmuseum, Stockholm.
Kandalakscha, 21-27 fm. Coll. N. Knipowitsch (from St. Petersbourg Museum).
Kandalakscha, 20-25 fm. Coll. A. Birula (from St. Petersbourg Museum).
Nordenskiold's Sea, 77 i 1 N., 114 35 1 E., 3.ix.i90i (Russian Polar Expedition,
1900-1903, St. 46).
Nordenskiold's Sea, 75 42 1 N., 124 41 1 E., 6.ix.i90i (Russian Polar Expedition,
1900-1903).
East Greenland, Franz- Josef Fjord (Muskoxfjord), 21.viii.1900, 220 m. (Swedish
zoological Polar Expedition, 1900, St. 27).
All the material from the above localities is sterile. The pseudohydrotheca of the
hydranth reaches almost to the tentacles, and in one hydranth it is a little loose-
fitting and wrinkled which permits its limits to be seen. There is no indication of a
web between the tentacles.
The most interesting (and also the characteristic) feature of this hydroid is the
formation of ' nodes ' on the stems, and if we accept the belief that the species is a
gymnoblastic hydroid (as seems most likely), these "nodes" indicate points of
regeneration after periodic dying down of the hydranths. The species is referred
provisionally to Bougainvillia.
SYNOPSIS OF THE STATUS OF THE DIFFERENT SPECIES
OF " PERIGONIMUS "
Order ATHECATA (ANTHOMEDUSAE)
Family Cytaeidae
Genus Perarella Stechow (Original name)
Perarella schneideri Perigonimus schneideri Motz-Kossowska
Family Bougainvillidae
Sub-family Thamnostominae
Genus Thamnostoma Haeckel
Thamnostoma cidaritis
Sub-family Bougainvillinae
Genus Bougainvillia Lesson
Bougainvillia macloviana
Bougainvillia muscoides
Bougainvillia ramosa
Bougainvillia linearis
Perigonimus cidaritis Weismann
Perigonimus macloviana Vanhoffen
Perigonimus muscoides M. Sars
Perigonimus muscus Allman
Atractylis linearis Alder
A REVISION OF THE HYDROID GENUS PERIGONIMUS M. SARS 347
Provisionally also:
Bougainvillia nanellus
Bougainvillia robusta
Bougainvillia vagans
Bougainvillia yoldiae-arcticae
Bougainvillia sp.
Genus Dicoryne Allman
Dicoryne conferta
Genus Rhizorhagium M. Sars
Rhizorhagium roseum
Rhizorhagium antarcticum
Rhizorhagium formosum
Perigonimus (?) nanellus Stechow
Perigonimus robustus Fraser
Perigonimus vagans Thornely
Perigonimus yoldiae-arcticae Birula
Atractylis quadritentaculata Wright
Eudendrium confertum Alder
Rhizorhagium roseum M. Sars
Perigonimus sarsii Bonnevie
? A tractylis coccinea Wright
? A tractylis miniata Wright
? Perigonimus inflatus Duerden
Perigonimus antarcticus Hickson & Gravely
Perigonimus formosus Fewkes
Family Pandeidae
Sub-family Stomotocinae
Genus Stomotoca L. Agassiz
Stomotoca dinema
Stomotoca georginae
Sub-family Pandeinae
Genus Leuckartiara Hartlaub
Leuckartiara octona
Provisionally also :
Leuckartiara abyssi
Genus Halitholus Hartlaub
Halitholus cirratus
Family Eudendriidae
Genus Eudendrium Ehrenberg
Eudendrium ramosum
Perigonimus serpens Allman
Perigonimus (?) nutans Hincks
Perigonimus apicatus Brooks
Perigonimus georginae Hadzi
Eudendrium pusillum Wright
Eudendrium sessile Wright
Eudendrium repens
A tractylis palliata Wright
Perigonimus minutus Allman
Perigonimus vestitus Allman
Perigonimus jonesii Osborn & Hargitt
Perigonimus pugetensis Heath
? Perigonimus gelatinosus Duerden
Perigonimus abyssi G. O. Sars
Perigonimus cirratus Hartlaub
Perigonimus multicornis Allman
Order THECATA (LEPTOMEDUSAE)
Family Haleciidae
Sub-family Campanopsinae
Perigonimus nudus Stechow
Family Proboscidactylidae
Genus Pochella Hartlaub
? Pochella polynema
Order LIMNOMEDUSAE
A tractylis bitentaculata Wright
348 A REVISION OF THE HYDROID GENUS PERIGONIMUS M. SARS
Incertae Sedis
Perigonimus decorans Schneider
Perigonimus napolitanus Hargitt
Perigonimus steinachi Jickeli
Perigonella sulphureus [Perigonimus sulphureus Chun)
These few species which I have been unable to classify are insufficiently described.
The above classification is in accord with recent progress, but some of it may not
be familiar to specialists who work only on hydroids and leave the medusa phase out
of consideration. I believe with Allman (1864a) that "An adequate conception of the
hydroid can thus only be obtained by regarding it as the product of two factors, one
of them finding its expression in the trophosome, and the other in the gonosome ".
The result is frequently a mosaic, a blending of characteristics into a pattern which
gives a much better picture of the position of the living species than does consideration
of only a part of the life history. 1
Much has been written by Kramp (1941 and 1949) about the dangers of attempting
a single classification for hydroids and medusae, but its outlines are already apparent
for the Anthomedusae, and this revision of Perigonimus is intended as a contribution
towards it.
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ON THREE NORTHERN SPECIES OF
HYDRACTINIA
By WILLIAM J. REES
(With Plates 11-12)
SYNOPSIS
Three species of Hydractinia, H. allmani Bonnevie, H. carica Bergh and H. monocarpa Allman
from the Arctic are redescribed from numerous colonies. The great differences noted between
colonies of the same species are shown to depend on the stage of reproductive exhaustion reached
when the colonies were taken. What is known of their biology and distribution is summarized.
INTRODUCTION
Hydractinia is a large genus, and, of the many species which have been described,
few have been worked on alive prior to a description being drawn up. In consequence
many of the specialists who have worked on these animals (perforce from Expedition
material) have seldom considered the specimens before them as having been something
alive and constantly changing. Inevitably phases in growth and reproduction as well
as ecological forms have on occasion been described as distinct species.
The northern Hydractinia species have been partially revised by Broch (1916) and
Kramp (1932) so that there is now less confusion about the identity of arctic and
subarctic forms. Some species however require redescription based on re-examination
of as many colonies as possible in order to eliminate the use of so-called diagnostic
features which are due to growth, seasonal variation, reproductive activities or
reflect environmental conditions.
In species like Coryne where the gonophores are borne directly on the body of the
hydranth, these reproductive activities cause the hydranth itself to be resorbed and
often reduced to a mere stump without tentacles. This phenomenon may be termed
reproductive exhaustion and it is by no means confined to the capitate hydroids
where its effects are most noticeable. In Hydractinia reproductive exhaustion mani-
fests itself first in the fertile polyps (or special reproductive polyps as the case may be)
and also causes some reduction in the size of the nutritive polyps. I first noticed an
appreciable reduction in the size of the sterile polyps in a Hydractinid due to this
cause in the undescribed hydroid of Podocoryne borealis. During the fifteen days I had
it under observation, it was producing large numbers of medusae, and by the time
I was certain that it was distinct from the better-known P. carnea, the colony had
altered so much in appearance that I could not honestly describe it in that condition
(Rees, 1941).
352 ON THREE NORTHERN SPECIES OF HYDRACTINIA
This paper is confined to observations on three northern species, Hydractinia
allmani Bonnevie, Hydractinia carica Bergh and Hydractinia monocarpa Allman, and
is based on the rich arctic and subarctic collections of the Naturhistoriska Riksmuseum,
Stockholm. Some of the material has been briefly described by Elof Jaderholm, but
all the locality records of colonies in the Riksmuseum have been reported by him in
various papers.
Hydractinia allmani Bonnevie
Hydractinia allmanii Bonnevie, 1898, Z. wiss. Zool., 63 : 485, pi. xxvi, figs. 36-37.
Bonnevie, 1899, Norske Nordhavs Exped., 1876-8, 26 : 47, pi. i, fig. 1.
Jaderholm, 1902, Bih. Svenska Vet. Akad. Handl. 28, Afd. 4, No. 12 : 8.
Broch, 1903, Bergens Mus. Arb. 1903, No. 9, Table.
Hydractinia allmani, Hartlaub, 1905, Zool. Jb. Suppl. 6, Bd. 3 : 518.
Jaderholm, 1908, Mem. Acad. Sci. St. Petersb. (8) 18, No. 12 : 7, pi. i, fig. 5 ; pi. ii, figs.
2-5-
Jaderholm, 1909, K. Svenska Vetensk-Akad. Handl. 45, No. 1 : 49.
Broch, 1910, Fauna Arctica, 5 : 200.
Kiihn, 1910, Zool. Jb. (Anat.), 30 : 107.
Kramp, 1914, Medd. Gronland, 23 : 988.
Broch, 1916, Danish Ingolf Exped. 5 : 44.
Rylov, 1923, Annu. Mus. Zool. Acad. St. Petersb., 24 : 151.
Kramp, 1943, Medd. Gronland, 121, No. 11:8.
Type locality. Norske Nordhavs Expedition 1876-78, St. 137, 67 24' N.,
8° 58' E., 827 m. (Zoological Museum, Oslo) Type locality selected here.
Material seen. In Riksmuseum, Stockholm :
Swedish Polar Expedition, 1900.
St. 17, Mackenzie Bay, north of Franz Josef Fjord, E. Greenland, 12-35 m -
St. 23, Mackenzie Bay, north of Franz Josef Fjord, E. Greenland, 3-10 m.
St. 27, Muskoxfjord, Franz Josef Fjord, E. Greenland, 220 m.
Russian Polar Expedition, 1 900-1 903.
St. 10 d, Kara Sea, 73 27' N., 79 15' E., 40 m.
St. 18, Gulf of Middendorff, 75 54' N., 92 59' E., 12-14 m.
St. 50, north of New Siberian Isles, 77 20' 30" N., 138 47' E., 38 m.
Otto Torell : Bellsund, Spitzbergen, 8-12 fm.
Specific Characters. Hydractinia with encrusting base, with few simple, smooth
spines. No tentaculozooids or spiral zooids. Nutritive polyps, long columnar
(1-4-15-0 mm. high) with up to 16 filiform tentacles. Reproductive polyps (075-5-0
mm.) at first with hypostome and at least 8 tentacles becoming reduced to stumps
with armed proboscis and even without tentacles. Usually one large and one small
gonophore on each reproductive polyp.
Gonophores (o-6 — 1-4 X 0-5 — 1-3 mm.). Female, cryptomedusoid with 4 well-
developed radial canals and 8 teat-like tentacle rudiments. Eggs numerous, polygonal
at first becoming rounded or oval and escaping through a pore between the tentacle
rudiments. Male gonophores also cryptomedusoid.
ON THREE NORTHERN SPECIES OF HYDRACTINIA 353
Distribution. Apart from two records in rather deep water off the Norwegian
coast, viz : position, 67 24' N., 8° 58' W., S.W. of the Lofotens at a depth of 827 m.
(Bonnevie, 1899) and position 63 12' N., i° 30' E., at a depth of 1320 m. (Broch,
1903), H. allmani has been found mainly in the high Arctic.
It has been found in the Spitzbergen area on a number of occasions (Bonnevie,
1899 ; Jaderholm, 1909 and Rylov, 1923). To the east the Russian Polar Expedition
1900-1903 took it in the Kara Sea, the Gulf of Middendorff and north of the New
Siberian Islands (Jaderholm, 1908). In Greenland it was taken at three stations by
the Swedish Polar Expedition, 1900, in the neighbourhood of Franz Josef Fjord
(Jaderholm, 1902) and in Kempes Fjord by the Swedish Gronland Expedition, 1899.
Other Greenland records from Hurry Fjord (Scoresby Sound area), Kempes Fjord,
Vega Sound and the mouth of Forsblad Fjord are given by Kramp (1943).
The absence of records from the Okhotsk Sea and the Bering Sea are possibly due
to lack of observations and the species may prove to be circumpolar in distribution.
Description. This is a large and handsome Hydractinia, but small and intermediate
colonies occur and the species is very variable. For this reason several colonies are
described below.
Fully grown colonies before the onset of reproductive exhaustion, as already
mentioned above, are large and handsome and the following description is based on a
colony from Muskoxfjord, East Greenland (PL 1 1 , figs. 3-4). The encrusting base, which
is continuous, has few spines, and arising from it are long spindly nutritive polyps
reaching a height (to hypostome) of 8-0-13-5 mm. in preserved material. There is a
well developed, conical proboscis, which is armed with a few nematocysts, with at
its base, a circle of about 15 tentacles in a close double whorl. Each hydranth arises
directly from the hydrorhiza and there is no distinct ring of perisarc at this point
although the perisarc forms a thin wrinkled sheath.
The largest reproductive hydranths are 1-6-4-5 mm. high, that is, they are only
about half the length of the nutritive ones, and from this colony it is not possible to
say whether they are derived from nutritive polyps. The largest seen had a length of
4-5 mm.; it had a conical proboscis moderately well armed with nematocysts at its
distal end. The tentacles were four in number and only 0-2 mm. in length. Some
distance below the head of the hydranth it carried one large and one small gonophore.
Other blastostyles ranged down to i-6 mm. in height and the tentacles were some-
times reduced to mere knobs. Each blast ostyle usually carried one large and one
rudimentary gonophore. None of the gonophores was fully developed, the largest
having a diameter of 1-4 mm. No spiral zooids have been observed.
Both male and female gonophores occur in the same colony. They are cryptomedu-
soids with well developed radial canals and the tentacle bulb rudiments are repre-
sented by four slightly pigmented areas (more mature gonophores are described from
the colony taken at Mackenzie Bay, East Greenland on 1st August, 1900).
Another colony from Mackenzie Bay (also East Greenland) has similar gonophores,
but the nutritive polyps are either less well developed or retracted (due to preserva-
tion methods) (PL 1 1 , figs. 1 and 2). The polyps have a height of up to 5 mm. Likewise
the gonophores are not fully mature and the blastostyles themselves are quite small
being only 1*3-1-5 mm. in total height.
354
ON THREE NORTHERN SPECIES OF HYDRACTINIA
A second colony from Mackenzie Bay taken on ist August, 1900, has distinctly
smaller hydranths, these being less than 3-0 mm. in height, but both male and female
gonophores are fully developed. The female gonophore is fully developed at 1-2 mm.
in diameter and the tentacular rudiments are represented by 4 - 8 teat-like processes
and the manubrium may protrude through the opening by which the eggs escape.
Another colony taken by the Russian Polar Expedition, 1900-1903, St. 50, north
of the New Siberian Islands, is remarkable for the length of the nutritive polyps
which are up to 14-3 mm. in height. The female gonophores are ripe (PL 11, fig. 7) and
the male ones nearly so. The blast ostyles vary from 1-5 mm. to 3-3 mm. in height
and the larger carry 8 tentacles and the smaller ones only 1 or 2. Typically only 2
gonophores, one large and one small, are found on each blastostyle. The ripe female
gonophore has 4 perradial rudiments of tentacles and 4 smaller interradial rudiments
(pi. 11, fig. 7 ).
The individual (both nutritive and reproductive) polyps of a colony from the Kara
Sea are small. Some of the female gonophores are sufficiently advanced to show the
rudiments of tentacle bulbs and radial canals. Here the blastostyles are more reduced
than in some of the other colonies and often only one or two stumps of tentacles are
left (PI. 11, figs. 5 and 6). The proboscis of the reduced blastostyle is well armed with
nematocysts. Eggs are numerous at first but become reduced in number to 10-15
in the ripe gonophore.
As will be noticed from Table I there is a considerable range in the size of the
nutritive polyps and the very large size of the polyps in two colonies (one from
Mackenzie Bay, St. 27 and the other from north of the New Siberian Islands) made
me suspect that a different species was involved, but a careful re-examination of the
material revealed no essential points of difference in any of the colonies except in size
and degree of maturity of the gonophores.
Table I. — Hyd? -actinia allmani Bonne vie.
(Measurements in mm.)
Length of nutritive polyp
Maximum diameter of polyp
Length of hypostome
Height of reproductive polyp
Length of gonophore
Diameter of gonophore .
Mackenzie Bay, E.
Greenland
Gulf of
North of
,^.
Middendorff,
New Siberian
c
*>
St. 17
St. 23
St. 27
Kara Sea
Islands
. 2-4-2-7
1 -4-3- 1
7-o-i3-5
2 • 0-2 • 2
3-1-12-0
. 0-4-0-5
• 2-0 • 4
0-5-0-9
• 2-0 • 7
• 7-0 • 9
. 0-5-0-6
0-3-0-4
• 6-0 • 8
0-3-0-35
• 5-0 • 8
• 1 -5-i -7
1 -1-2-9
2-3-4-5
0-75-1-4
1-5-3-3
. 1-1-1-3
• 9-1 • I
o-6-i -4
• 6-1 • 1
1 -i-i -4
. I-Q-I-2
• 9-1 • I
0-3-1-3
o-5-i-i
1-0-1-3
In these colonies the nutritive polyps are more than twice the size of the largest
reproductive polyps ; the latter have up to 8 tentacles and are very similar to the
sterile polyps. It is not possible to be completely certain that they are reduced from
the nutritive polyps through bearing gonophores, as in Podocoryne carnea, but it may
be that we have here the beginnings of the specialization which results in the pro-
duction of the specialized reduced reproductive polyps seen in Hydractinia echinata.
ON THREE NORTHERN SPECIES OF HYDRACTINIA 355
In the remaining colonies there is some overlap in the size of the two kinds of polyps,
and in view of the small size of all the polyps it appears likely that they are reduced
by prolonged breeding. This raises the question of the status of Hydr actinia ornata
Bonne vie, which, as far as I can see, is distinguished from H. allmani only on size ;
its status must await the re-investigation of the original material.
There is one feature in the reduction of the reproductive polyp which is worth
noting. The proboscis of the very reduced polyp is armed with nematocysts, these
being concentrated at its tip as the tentacles are resorbed. The use of this character
(an armed proboscis) in classification should therefore be used with caution.
Biology. In all records where the substratum of Hydr actinia allmani has been
indicated, the gastropod involved has been Sipho curtus (Jeffreys). In the high Arctic
the hydroid occurs at depths of 3-760 m. in Greenland, 75-350 m. off Spitzbergen
and in 12-40 m. in the more easterly localities.
Records of this species from off the Norwegian coast in comparatively deep water
(see Distribution) suggest that this submergence is related to temperature. A colony
reported by Jaderholm (1909, p. 49) was dredged from 350 m. where the bottom
temperature was 273 C, while Bonnevie (1899, p. 11) recorded the species from
bottoms where the temperatures were — i° C. and — 1-2° C. Comparable tempera-
tures (given by Ekman, 1953, p. 166) exist at the other localities at which this
species has been found.
It is probable that Hydractinia allmani can be regarded as a stenothermal cold
water species.
Historical and Relationships. Since Bonnevie described the species in
1898, numerous records indicate that this is a common species in the high Arctic.
It can be readily distinguished from Hydractinia monocarpa and H. echinata by its
simple smooth spines ; from Podocoryne it is separated by its fixed gonophores.
Compared with H. carica, it is more robust in habit, with longer polyps and more
polyp-like reproductive zooids. The numerous eggs in the female gonophores (eggs
are few in number in the gonophores of H. carica) also help to distinguish it.
Hydractinia carica Bergh 1887
Hydractinia carica Bergh, 1887, Lutken, C.F. Dijmphna-Togtets zool.-bot. Ubbytte, p. 331, pi.
xxviii, fig. 1.
Schydlowsky, 1902, Trav. Soc. Nat. Univ. Kharkow, 36 : 114.
Hartlaub, 1905, Zool. Jb. Suppl., 6, Bd. 3 : 518.
Jaderholm, 1908, Mem. Acad. Sci. St. Petersb. (8) 18, No. 12:8.
Jaderholm, 1909, K. Svenska Vetens.-Akad. Handl., 45, No. 1 : p. 48, pi. ii, figs. 10 and 11.
Broch, 1 910, Fauna Arctica, 5 : 200.
Broch, 1916, Danish I ngolf Expedition, 5, Part 6 : 48, pi. i, fig. 11.
Scheuring, 1922, Wiss. Meeresuntersuch. Abt. Helgoland, 13 : p. 168.
Rylov, 1923, Annu. Mus. Zool. Acad. St. Peterb., 24 : 150, Taf. vi, figs. 2-3.
Fraser, 1931, Contr. Canad. Biol., n.s., 6 : 6.
Uschakow, 1937, Trans. A ret. Inst. Leningr. 50 : 12.
Fraser, 1944, Hydroids Atlant. Coast N. America, p. 77, pi. xiii, fig. 53.
356 ON THREE NORTHERN SPECIES OF HYDRACTINIA
Hydractinia minuta Bonnevie, 1898, Z. wiss. Z00L, 63 : 468, pi. xxvi, fig. 38.
Bonnevie, 1899, Norske Nordhavs Exped. 1876-8, 26, Zool. : 47 pi. i, fig. 3.
Bonnevie, 1901, Meeresfauna von Bergen. Hydv., 1 : 7.
Broch, 1910, Fauna Arctica, 5 : p. 141 and 200.
Type locality. Petuchoffskoi Schar, Kara Sea, 15 m. (Dijmphna Expedition —
Zoological Museum, Copenhagen) .
Material seen. In Riksmuseum, Stockholm :
Fragment of type material on a mounted slide.
Bellsund, Spitzbergen, 20 fm. 22nd July, 1864 (A. J. Malmgren)
Bellsund, Spitzbergen, 8-12 fm. (Otto Torrell)
Recherche Bay, Spitzbergen, 13th July, 1898 (Spitzbergen Expedition, 1898)
67 56' N., 66° 18' W., 13th October, 1897 (E. Nilsson)
Russian Polar Expedition, 1900-1903, St. 3, Murman Sea, 69 35' N., 50 5'
E.; 70 m.
In British Museum :
Klaas Billen Bay, Spitzbergen, 28th July, 1921, C. S. Elton (Oxford Univ.
Exped. to Spitzbergen, 1921).
Specific characters. Hydractinia with encrusting base with few high conical
smooth spines (up to 07 mm. high) with rounded apices. Nutritive polyps, small, up
to 2-3 mm. high when fixed, with a single whorl of 10-16 tentacles. Proboscis simple,
almost unarmed. No spiral zooids known.
Reproductive polyps up to 2-3 mm. high, at first with conical hypostome and at
least 8 tentacles, becoming greatly reduced to very short stumps without tentacles ;
proboscis armed with nematocysts when present. Gonophores 3-7 in number borne
in a ring round the polyp.
Gonophores crypt omedusoid, with 4 radial canals and rudiments of tentacles
present but not developed into bulbs. Eggs large, 4-10 in number.
Distribution. Hydractinia carica is known from the Kara Sea (Bergh, 1887),
the Barents Sea (Jaderholm, 1908 ; Scheuring, 1922), Spitzbergen, the Davis Strait
area (West Greenland) and on the Norwegian coast in the neighbourhood of Bergen.
There are numerous records of this species from Spitzbergen, most of these being
summarized by Jaderholm (1909) and Rylov (1923). I have been able to verify
Jaderholm's record (1908) from Davis Strait and there are others by Fraser (1931 and
1944 as carica and minuta) from approximately the same area. The most southerly
record is that of Bonnevie (1901) from the Hjeltefjord, Bergen as H. minuta.
There is insufficient information to hazard any opinion as to whether this species
has a circumpolar distribution.
Description. This is an encrusting species with large spines but without small
prickles. The large spines are simple, smooth and not sharply pointed ; they have no
pore at the tip and in size range up to 07 mm. in length.
In preserved material the hydranths are tubular to fusiform in shape and may be
over 2-o mm. in height. The head of the hydranth is sometimes clavate and there is a
ON THREE NORTHERN SPECIES OF HYDRACTINIA
357
conical proboscis without many nematocysts around the mouth. There are usually
about 16 tentacles arranged more or less in a single whorl.
The reproductive polyps vary considerably from colony to colony according to the
stage they had reached when taken, and it has been deemed desirable to describe
them in the different colonies.
In the colony from Bellsund, Spitzbergen the reproductive polyps are completely
reduced to very short stumps 0-17 mm. or less in length, bearing up to 5 gonophores
each. These have diameters of about 0-45 mm. Sterile hydranths are 1-3-2-0 mm. in
length.
Another colony from Recherche Bay, Spitzbergen, does not appear to have reached
the same state of reproductive exhaustion as the one from Bellsund. In this colony
the fertile polyps are still recognizable as polyps ; one of these still possessed 8 fully
developed tentacles and a distinct proboscis and carried 5 moderately developed
gonophores. Another reproductive polyp, with a height of 0-9 mm., had 7-8 tentacles
and 4 gonophores each with a diameter of 0-3 mm.
Text-fig. 1. — Hydractinia carica Bergh : blastostyles from a colony from Recherche
Bay, Spitzbergen (Swedish Spitzbergen Expedition, 1898).
A colony from position 67 56' N., 66° 18' W., had particularly large blastostyles,
1-0-2-3 mm - i n height to hypostome. These had 3-5 gonophores each and only a few
had up to 2 reduced tentacles. The hypostome was usually well armed with nemato-
cysts.
There were only a few very reduced polyps. These were so short that they could
not be measured accurately ; they carried 5-7 gonophores and were clearly reaching
the exhaustion stage.
Colonies from the Murmansk coast and Bellsund, Spitzbergen, were fertile and some
of the reproductive polyps of these possessed 2 or more tentacles.
A mounted slide marked Orig. Ex., Dijmphna Exp. presented by Levinsen appears
to consist of part of the type material of this species. Measurements of this and other
colonies in the Riksmuseum are given in Table II. Some of the reproductive polyps
carry about 4 tentacles and others have lost them (Text-fig. 1).
358
ON THREE NORTHERN SPECIES OF HYDRACTINIA
Table
II,
— Hydractinia carica Bergh
(Measurements in mm.)
Kara Sea
(fragment of
67 56' N.,
Bell Sound,
type material)
66° 18' W.
Spitzbergen
Length of polyp
1-5-1 -7
1 • 0-2 • 3
1 • 0-2 •
Maximum diameter of polyp
0-3-0-4
• 1-0 • 2
• 2-0 • 3
Length of hypostome
o- 4
0-2
• 1-0 • 3
Height of reproductive polyp .
o • 75-0 • 9
0-1-2-3
■ 1 7-0 • 2
Length of gonophore
o • 28-0 • 4
0-3-0-4
o-45
Diameter of gonophore
• 25-0 • 4
0-3-0-4
o-45
Length of spine
—
—
0-37-0-7
Only female gonophores have been noted in this material ; they are cryptomedu-
soids without radial canals but with apical patches of nematocysts. There is a well-
developed manubrium and when ripe the gonophores contain 4-10 large eggs each
(Text-fig. 2, a, b).
Text-fig. 2. — Hydractinia carica Bergh ; gonophores from a fragment of the type colony,
Kara Sea (Dijmphna Expedition) : a, female gonophore ; b, another female gonophore
in optical section.
This material agrees with the earlier descriptions of Bergh (1887), Jaderholm
(1908) and Broch (1916), but it has been possible to get a better picture of the species
as a whole. The nutritive polyps (in preserved material) are seldom more than 2-0
mm. in height and it is possible that the reproductive polyps are derived directly
from them. Bergh and the other authors indicated that the reproductive polyps
might have 1-4 tentacles, but I have found 8 which supports my suggestion. Jader-
holm (1909, pi. ii, figs. 10 and n) gives good illustrations of partially reduced hydranths,
and others are figured here (Fig. 1), but they may become so short that even the
hypostome is almost completely reduced and the stalk below the point of origin of
the gonophores also becomes extremely short. Broch (1 916, p. 48) is undoubtedly
right in his contention that Bonnevie's H. minuta (Bonnevie, 1899, p. 48) is identical
with H. carica. I regard the colony she described as being in an advanced stage of
reproductive exhaustion.
Biology. This species favours the shells of various northern species of Buccinum
as a substratum. In the Barents Sea it has been found on Buccinum undatum and
B. tenue. In the Spitzbergen area Jaderholm (1909) has found it on Buccinum
glaciate and Trophon clathrus, while Rylov (1923) reported it as H. minuta on Buccinum
ovum and B. ciliatum.
ON THREE NORTHERN SPECIES OF HYDRACTINIA 359
The bathymetric range appears to be 0-120 m. in the Spitzbergen area and 40-70
m. in the Barents Sea.
Hydractinia carica may be regarded as mainly a cold water species which, if
Bonnevie's record from Bergen is authentic, is probably less stenothermal than H.
allmani.
Historical and Relationships. Hydractinia carica was described by Bergh
(1887) from Petuchoffskoi Schar in the Kara Sea from a depth of 15 metres.
Subsequently it was recorded many times in the far north and as H. minuta Bonne vie
from Spitzbergen and elsewhere. As a species it is distinguished from other northern
Hydractinia by its simple smooth spines, the extreme reduction which takes place
in its reproductive polyps and by its gonophores. These have no radial canals and
the female ones have few eggs.
Hydractinia monocarpa Allman
Hydractinia monocarpa Allman, 1874, Nature, Lond. 11, No. 270 : 179.
Marktanner, 1895, Zool. Jb. (Syst.), 8 : 394.
Schydlowsky, 1902, Trav. Soc. nat. Univ. Kharkow, 36 : 114.
Hartlaub, 1905, Zool. Jb. Suppl. 6, Bd. 3 : 518. '
Jaderholm, 1908, Mem. Acad. Sci. St. Petersb. (8), CI. Phys.-Math. 18, No. 12 : 8, Taf. 1,
fig. 6, Taf. 2, figs. 6-9.
Jaderholm, 1909, K. Svenska Vetens.-Akad. Handl. 45, No. 1 : 49, Taf. ii, figs. 12-13.
Broch, 1910, Fauna Arctica, 5 : 141.
Broch, 1916, Danish Ingolf Exped. 5, Pt. 6 : 47.
Rylov, 1923, Annu. Mus. Zool. Acad. St. Petersb. 24 : 148, Taf. VI, fig. 7.
Kramp, 1932, Medd. Gronland, 79, No. 1 : 16.
Uschakow, 1937, Trans. Arct. Inst. Leningr., 50 : 12.
Fraser, 1944, Hydroids Atlantic Coast N. America, p. 77.
Hydractinia echinata, Wintrier, 1880, Naturh. Tidsskr. 12 : 227, 254.
Levinsen, 1893, Vidensk. Medd. naturh. Foren Kobenhavn, 1892, p. 153.
Type locality. Spitzbergen, on Trophon clathratus L. (Allman, 1876) (Zoological
Museum, Copenhagen)
Material seen. In Riksmuseum, Stockholm :
Matotschkim Schar, 15 fm. (Novaya Zemlya Expedition, 1875).
Russian Polar Expedition, 1900-1903, St. 53, 77 10' N., 142 48' E. (N. of New
Siberian Isles) 35 m., on Trophon clathratus and Beta plicij era.
Specific characters. Hydractinia with encrusting base with long, often irregular,
hollow spines which may bifurcate distally. Spines longitudinally keeled especially
at base, but smooth and not thorned, up to 2-4 mm. high. Nutritive polyps fusiform
to tubular, up to 3-6 mm. high, borne on the encrusting base or on the sides of the
spines. Tentacles, 10-15. Spiral zooids not known.
Reproductive polyps, up to 1-5 mm. high, with armed proboscis and at least 4
tentacles, or may be reduced without tentacles and greatly shortened. Gonophores
1-4 on each, but usually only one.
360 ON THREE NORTHERN SPECIES OF HYDRACTINIA
Gonophores : female, up to 1-4 mm. in diameter ; cryptomedusoid with 4 radial
canals and apical patch of tissue. Male gonophores similar, up to 0-55-0-6 mm. in
diameter.
Distribution. All records of this species are from the high Arctic, chiefly from
Spitzbergen, Novaya Zemlya and the New Siberian Isles. It appears to be a common
species on shells at Spitzbergen (Airman, 1876 ; Jaderholm, 1909 and 1916 ; and
Broch, 1910). The Russian Polar Expedition 1900-1903 took it at two stations off
the New Siberian Islands and in Chatanga Bay in Nordenskj old's Sea (Jaderholm,
1908). Records from Novaya Zemlya are given by Jaderholm (1909).
West Greenland records from Upernivik and from near Cape Atholl are given by
Kramp (1932). This distribution suggests that the species is circumpolar.
Description. This species has an encrusting base, but the most characteristic
feature is the presence of long, upright irregularly-formed spines rising to a height of
about 3 mm. (PL 12, figs. 8-1 1). The spines are seldom straight or symmetrical and
are usually curved towards the tip which tapers to an open hyaline or horn-coloured
tube. Frequently the spines are double or bifurcated and their surface is seldom
smooth. The naked coenosarc may cover a good part of the spines.
The nutritive hydranths may be long and tubular reaching (when expanded) a
considerable length. Some preserved examples, moderately well fixed, have a length
of 3-6 mm., but, when contracted, may only be one- third or less than this length.
Such retracted polyps are either cylindrical or flask-shaped (PL 12, figs. 9 and 10).
The hypostome carries nematocysts, particularly near the tip, but they are few in
number. There are 10-14 tentacles. A curious feature of this species is that one or
more polyps may be carried on the spines themselves and I have seen up to three on
one spine (PL 12, figs. 9 and 10).
The reproductive polyps, noted in the material seen, were up to 1-5 mm. in height
and were provided with up to 4 tentacles. Each had an armed tip to the proboscis
and carried up to 4 gonophores. Only female gonophores have been noted by me ;
these are large, 0-5-0-7 mm. in diameter and are cryptomedusoids with manubrium
and radial canals. It is clear that the blast ostyles become considerably reduced due
to reproductive exhaustion. Frequently they are reduced so much that the gonophoi cs
appears to be almost sessile on the coenosarc. When this happens the upper part cf
the polyp seems to become larger and more polypoid, but the stalk below the point
of origin of the gonophore is extremely short.
In his original description Allman (1874) described the reproductive polyps as
devoid of tentacles, but tentacles have been noted by Jaderholm (1908). Rylov
(1923), Kramp (1932) and by myself. The diagnosis of the species (p. 25) differs
greatly from that of Allman because more colonies have been available for study.
Biology. Hydradinia monocarpa has been found on the shells of gastropods such as
Trophon clathrus, Beta plicifera and Buccinum hydrophanum (Jaderholm, 1908, 1916,
and Kramp, 1932). The hydroid may grow all over the shell as in a specimen of
Beta plicifera from Kol Bay, Isfjord, Spitzbergen. The nutritive and reproductive
polyps flourish best in and near the sutures between whorls, but they also occur on
exposed surfaces and are absent only on abraded surfaces of the shell. The tall spines
are fairly closely grouped and must offer some protection to the zooids.
ON THREE NORTHERN SPECIES OF HYDRACTINIA 361
The species has a known bathymetric range of 14-165 metres, the deepest record
being from West Greenland (Kramp, 1932). Most of the records from Spitzbergen
and elsewhere in the Polar Sea are, however, from shallow coastal waters.
Hydractinia monocarpa may be regarded as a high Arctic or stenothermal cold
water species.
Historical and Relationships. Hydractinia monocarpa was described by
Allman from a colony labelled " Spitzbergen " in the Zoologiske Museum, Copen-
hagen. It has since been found frequently in the Spitzbergen area and elsewhere in
the high Arctic.
This distinctive species, with its long, hollow, often bifurcated spines, its polyps
arising from both encrusting base and spines, its cryptomedusoid gonophores, with
four radial canals but without tentacle rudiments, cannot be mistaken for any other
northern species.
The presence of hydranths on the spines recalls the much greater development of
this feature in Hydrissa sodalis (Stimpson) from East Asiatic seas.
ACKNOWLEDGEMENTS
I wish to thank Dr. Bengt Hubendick for hospitality while examining this material
at the Riksmuseum, Stockholm, and also for the loan of some of it for more detailed
examination in London. I also wish to thank Mr. M. G. Sawyers for the care he has
taken in preparing the photographs of Hydractinia allmani and H. monocarpa from
slides made by the late Elof Jaderholm.
discussion
Colonies of Hydractinia allmani show marked variation according to the stage
of reproductive exhaustion reached at time of capture and it has been suggested here
that the reproductive polyps in this species are derived from nutritive ones. The
great variation in the size of nutritive polyps and in the reproductive polyps (from
well formed ones to almost blastostyles) is highly significant, and implies that we
must use these features with great caution in distinguishing species. It brings to mind
the very large number of boreal and tropical species of Hydractinia which have been
described on single colonies. A critical study of these must result in a great reduction
in the number of recognizable species. Another feature, noted in this paper, is the
concentration of nematocysts in the tip of the hypostome as the tentacles are resorbed
during reduction of the polyp. It means that the armed proboscis, too, must be used
with discretion as a diagnostic character.
The study of Hydractinia carica also bears out the general conclusions noted
above, but this species is notable for the almost complete resorption of the repro-
ductive polyp and shortening of its stem, so that the gonophores sometimes give the
appearance of being sessile on the hydrorhiza and even the hypostome may disappear.
This, too, is a factor to be considered in classification.
Although H. monocarpa can be readily distinguished by its spines, its reproductive
polyps vary according to phase and bear out the general conclusions reached for the
other two species.
362 ON THREE NORTHERN SPECIES OF HYDRACTINIA
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Zool. Jb. Suppl. 6, Bd. 3 : 497-714, 142 text-figs.
Jaderholm, E. 1902. Die Hydroiden der Schwedischen Zoologischen Polar-expedition 1900.
Bih. K. Svenska Vet.-Akad. Handl. 28, Afd. 4, No. 12 : 1-11, Taf. 1.
1908. Die Hydroiden des Sibirischen Eismeeres, gesammelt von der Russischen Polar-
Expedition 1900-1903. Mem. Acad. Sci. St. Petersb. (8), CI. Phys.-Math. 18, No. 12 :
1-28, pi. i-iii.
1909. Northern and Arctic Invertebrates in the collection of the Swedish State Museum
(Riksmuseum) . IV. Hydroiden. K. Svenska Vetens.-Akad., Handl. 45, No. 1 : 1-124,
pi. i-xii.
1916. Die Hydroiden des Eisfjords. K. Svenska Vetens.-Akad. Handl. 54, No. 4
1-14, 1 text-fig.
Kramp, P. L. 1914. Hydroider. Conspectus Faunae Groenlandicae. Medd. Gvenland,
23 : 383-456.
1932. The Godthaab Expedition 1928. Hydroids. Medd. Gronland, 79, No. 1 :
1-86, 34 text-figs.
1943- The Zoology of East Greenland. Hydroida. Medd. Grenland, 121, No. 11 :
1-52, 4 text-figs.
Levinsen, G. M. R. 1893. Meduser, Ctenophorer og Hydroider fra Gronlands Vestkyst.
Vidensk. Medd. naturh. Foren Kbh. (5), 4 : 143-212, pi. 5-8.
Marktanner-Turneretscher, G. 1895. Hydroiden. Zool. Jb. (Syst.) 8 : 391-438, pi.
11-13-
Rees, W. J. 1 94 1. On the life history and developmental stages of the medusa Podocoryne
borealis. J. Mar. biol. Ass. U.K., 25: 129-141.
Rylov, V. M. 1923. Zoologische Ergebnisse der Russichen Expeditionen nach Spitzbergen.
Hydroid-Athecata. Annu. Mus. Zool. Acad. St. Peterb. 24 : 140-160, pi. vi.
Scheuring, L. 1922. Untersuchungsfahrt des Reichsforschungsdampfers " Poseidon " in
das Barentsmeer im Juni and Juli 1913. Wiss. Meeresuntersuch Abt. Helgoland, N.F. 13 :
159-183, Taf. V and text-figs. A-C.
Schydlowsky, A. 1902. Materiaux relatifs a la faune des polypes Hydraires des mers arc-
tiques. I. Les Hydraires de la Mer Blanche le long du littoral des lies Solowetsky. Trav.
Soc. Nat. Univ. Kharkow, 36 : 1-276, Taf. I-V.
Uschakow, P. 1937. Materials on the Hydroids of the Arctic Seas of the U.S.S.R. Trans.
Arct. Inst. Leningr. 50 : 5-34, 4 text-figs.
Winther, G. 1880. Fortegnelse over de i Denmark og dets nordlige Bilande fundne Hydroide.
Naturh. Tidsskr. 12 : 223-278.
PLATE ii
Hydractinia allmani Bonne vie
Figs, i and 2. — Fragments of a colony from St. 23, Mackenzie Bay, East Greenland (Swedish
Zoological Polar Expedition, 1900) ; x 10.
Figs. 3 and 4. — Fragments of a colony from St. 27, Muskoxfjord, Franz Josef Fjord (Swedish
Zoological Polar Expedition, 1900) ; x 5.
Figs. 5 and 6. — Reproductive zooids with female gonophores ; St. 18, Karahavet, Midden-
dorff's Sea (Russian Polar Expedition, 1 900-1 903) ; X 25.
Fig. 7. — Reproductive zooid with ripe female gonophore ; St. 50, north of New Siberian
Islands (Russian Polar Expedition, 1900-1903) ; x 14.
Bull. B.M. (N.H.) Zool. 3, 8.
PLATE 1 1 ,
Hvdr actinia allmani Born
PLATE 12
Hydr actinia monocavpa Allman from St. 53, north of New Siberian Islands (Russian Polar
Expedition, 1900-1903).
Fig. 8. — Spines and female gonophores ; X 12.
Figs. 9-11. — Spines and nutritive polyps; x 20, x 21, X 25.
Bull. 11. M. (A\M.) Zool. 3, 8.
PLATE I
11
10
Hydractinia monocarpa Allman,
r 2 MAR 1956
PRINTED IN GREAT BRITAIN BY
ADLARD AND SON, LIMITED,
B A RTHOLOMEW PRESS. DORKING
2 MAR 1956
THE LIZARD GENUS
APRASIA
H. W. PARKER
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 3 No. 9
LONDON: 1956
THE LIZARD GENUS
APRASIA ;
ITS TAXONOMY AND TEMPERATURE-
CORRELATED VARIATION
BY
H. W. PARKER
Pp- 363-385 ; 7 Text-figures
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 3 No. 9
LONDON: 1956
THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, is
issued in five series corresponding to the Departments
of the Museum, and an Historical Series.
Parts will appear at irregular intervals as they become
ready. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.
This paper is Vol. 3, No. 9 of the Zoological series.
PRINTED BY ORDER OF THE TRUSTEES OF
THE BRITISH MUSEUM
Issued February, 1956 Price Six Shillings
THE LIZARD GENUS
APRASIA ;
ITS TAXONOMY AND TEMPERATURE-
CORRELATED VARIATION
By H. W. PARKER
SYNOPSIS
The genus Ophioseps cannot be maintained ; its past recognition was due to errors of identi-
fication and observation. Aprasia is the most completely adapted of the Pygopodidae for
fossorial life, the family exhibiting a continuous series in degeneration of limbs and girdles, with
the development of a serpentiform habitus and cranial modifications for burrowing. There
are three species with a combined range from Victoria to the Monte Bello Islands but which
occur together in only a restricted area near Albany. Two of the species show a considerable
range of variation in the numbers of presacral vertebrae and scales. This variation appears
to be correlated with differing summer temperatures. Since the characters are meristic and
such characters have been shown, in other cold-blooded animals, to be directly affected by the
temperature prevailing during embryological development, it is suggested that the variation
may prove to be thermoplastic. The other taxonomically useful, but non-meristic, characters
show no significant variation in different climatic regions. The temperature correlated clines
are discontinuous geographically, permitting recognition of two new subspecies.
Two small limbless lizards were collected on Hermite Island, one of the Monte
Bello group, in 1952. They differ from any of the known species of the genus Aprasia
but most nearly resemble Aprasia repens Fry. Since the Monte Bello Islands are
geographically far removed from the recorded range of any species of Aprasia a
re-survey of the known species of the genus, and its alleged ally Ophioseps, was
necessary to ascertain the probable significance of the differences. More than 120
specimens were assembled for comparison, and this series covered not only the
whole of the previously known geographical range, but included recently captured
specimens from regions where none of these lizards had previously been reported.
MORPHOLOGY AND COMPOSITION OF THE GENUS
As will be demonstrated presently the series is subdivisible into five groups with
morphological differences, but all the specimens share the following characters :
Scales smooth, subequal dorsally and ventrally except on the throat and pectoral
region, where they are slightly smaller than on the belly ; 12 or 14 longitudinal
rows of scales at mid-body. General habitus worm-like, the head and tail not being
wider or narrower than the body, which is very elongate and cylindrical. The
snout projects in front of the labial margin and the large rostral shield is visible
ZOOL. 3, 9. 24
366
THE LIZARD GENUS APRASIA
from above ; five supralabials, the first being wholly or partly fused with a large
nasal shield, which forms a suture with its fellow behind the rostral ; pre-frontals
large, forming a median suture and in contact with the second labial ; a small
pre-ocular in contact with the third labial and the supra-ocular, which reaches the
fourth supra-labial or is separated from it by a single small post-ocular ; frontal
hexagonal ; parietals not much larger than the scales of the occiput and nape, not
in contact behind the frontal. Eye covered by a circular brill edged with a ring
of small granules. Shields and scales of the head beset with numerous minute
pustules, presumably tactile sense-organs. No external auditory meatus, tympanum
or tympanic cavity. No vestiges of fore-limbs externally ; hind-limbs visible as
small triangular flaps at the lateral corners of the vent, each being covered by a
single triangular scale not very different in size or appearance from the other scales
of the region. No pre-anal pores. A pair of post-anal sacs close behind the vent
and opening by a pair of small pores just within its posterior border. Bordering the
anterior margin of each of these sacs there is, in males, a post-cloacal bone which
pierces the skin laterally close, but caudomesial, to the hind-limb vestige ; the
protruding portion is sometimes claw-shaped, with the free terminal part of the claw
directed postero-dorsally, or sometimes subcircular in surface view (Text-fig. i).
The skull of Aprasia repens has recently been figured by McDowell & Bogert
(1954, fig. 21) and specimens of all the species represented in the present material
(vide infra) have been found to conform to this pattern. There is a single pre-
Fig. 1. Pelvic region of a cleared specimen seen in three-quarters ventro-lateral view
from the right side to show hind limb, girdle and post-cloacal bones. (Aprasia striolata,
THE LIZARD GENUS APRASIA 367
maxilla ; paired nasals form a long median suture ; the prefrontals are separated
mesially and extend posteriorly above the orbit to form a suture with a small post-
frontal ; the frontals are fused to form a single median element, but the parietals
are paired, forming a long median suture ; there is a narrow supra temporal
(tabular). 1 In both sexes the maxillary is edentulous and the dentary is toothed ;
the premaxilla is toothed in adult males of all species, but is completely edentulous
in juveniles of all species and also in females, except in A. striolata where vestigial
teeth may be present. In the lower jaw the splenial is wanting or is fused with the
prearticular. The only skull examined that deviates in any way from this pattern
is the one figured and described by Jensen (1901) as Ophiopsiseps nasutus Bocage.
This description has been the cause of much confusion. The lizard was incorrectly
identified and the description is incorrect in many points that subsequent authors
have accepted as indicating the existence of a genus (Ophioseps or Ophiopsiseps)
distinct from Aprasia. As will be demonstrated presently (p. 380) , the type specimens
of the type species of Aprasia Gray and Ophioseps Bocage are conspecific and the
two names are thus strict synonyms. But even apart from this nomenclatorial
consideration there appear to be no grounds for the recognition of a second genus.
Jensen's specimens have been compared with a series of others from the same
geographical area (Victoria and S. Australia) and have been found to be juveniles
of the one and only species that occurs there. This, in its adult phase, has a skull
like that described above. Jensen was in error in several respects, notably in
regard to the frontals, prefrontals, postfrontals and " squamosal " (supratemporal)
which, despite the immaturity of the skull, do not differ from those of the adult.
He was also misled by the absence of any median suture between the parietals
into believing that these bones were completely fused ; in fact, owing to immaturity,
the development of the parietals is incomplete, but they exist as two very thin
lateral blades widely separated on the mid-line.
Although all the species and subspecies examined have skulls that are essentially
similar in their general structure, there will undoubtedly be found to be specific
differences and, perhaps, ontogenetic differences in proportion. Insufficient skeletal
material is available to the writer to permit of any definite conclusions, but in the
long-snouted species (A. repens) the pre-orbital increase in length is accompanied
by an increase in length of the nasals. Thus, the ratio of frontal to nasal length
was found to be approximately 1-24 (c.f. also Bogert & McDowell's figure where the
ratio is 1-25) in this species, but 1-40-1-49 in specimens from South Australia
{A. striolata) and 1-50-1-56 in an example of Aprasia pulchella and the holotype of
Ophioseps nasutus Bocage.
Other skeletal features of the series are as follows. Presacral vertebrae 88-137,
the number varying from species to species ; the first three vertebrae (four in one
specimen) lack ribs ; sacral vertebrae three, with sacral ribs diminishing in length
caudad, the anterior two being free and the last knob-like and fused to the centrum
(Text-fig. 2). Shoulder girdle a V-shaped structure composed of a pair of rod-like
1 McDowell & Bogert (1954) figure this bone in their dorsal and ventral views (figs. 21 a and b) but
do not label it ; it is not shown in the lateral view (fig. 21c).
zool. 3, 9, 24§
368 THE LIZARD GENUS APHASIA
coracoids connected across the mid- ventral line by a V-shaped sternal cartilage.
Each coracoid, which is slightly expanded at its ends, runs antero-dorsally from
the sternum to a point near the distal extremity of the first rib. The dorsal end
of each is tipped with a short, tapering length of cartilage which may represent
a scapula. There are no traces of any bones of the fore-limb.
The pelvis (Text-figs, i and 2) consists of a subtriangular, slightly curled ilium
and an ischiopubis. The ilium articulates at its postero-dorsal corner with the
Fig. 2. Aprasia striolata striolata. Left pelvis and hind limb of a male, diagrammatic
to show sacral connexions.
third, knob-like, sacral rib and the other two sacral ribs have loose ligamentous
connexions with its antero-dorsal border. The ischium and pubis are fused, the
ischiadic portion of the combined bone being rectangular and the pubic portion
rod-like ; both portions are widely separated from their fellows on the mid-ventral
line. The hind limb consists of a short, stout femur, with a well-developed trochanter,
and a nodule-like vestige of a tibia ; sometimes there is also a vestigial fibula.
POSITION IN THE FAMILY PYGOPODIDAE
The various forms possessing these characters in common are clearly a closely
related group and this position is best expressed taxonomically by treating them
as a single genus. McDowell & Bogert (1954) in discussing the relationships of the
family Pygopodidae have pointed out that " the more typical pygopods, the Pygo-
podinae and Lialinae " are akin to the geckoes in many features. But Jensen's
account of Ophioseps misled them and, although they do not commit themselves
definitely, they say " we see no reason to question Jensen's placing of Ophioseps
in a distinct family, as Boulenger (1885) suggested. It is probable, however, that
Aprasia should be transferred to the Ophiosepidae." Since Ophioseps, sensu
Jensen, is a fallacy any such disposal of Aprasia would be absurd and it remains
THE LIZARD GENUS APRASIA 369
to consider its position in relation to the other genera of the Pygopodidae. It
shares with Pygopus, Paradelma, Delma and Lialis a unique character, a pair of
post-cloacal bones forming external hooks as a secondary sex character of the male.
These structures were first described and figured in 1870 by Fiirbringer in Lialis
(burtoni) but without the realization that they were associated with sex ; the same
author also described the limbs and girdles of Pygopus, but found no such structures
because, presumably, his specimen was a female. Boulenger (1885 : 239) mentioned,
as a family character, the presence of a spur laterally behind the vent, but wrongly
described it as a projection of the ischium and did not, apparently, observe that it
was only present in the one sex. Kinghorn (1923 : 134), alone, seems to have
observed that in Aprasia the spurs are present in males only. The present writer
has confirmed that they are, likewise, secondary sex characters of the male in the
other four genera mentioned above.
Post-cloacal bones occur in male geckoes (Smith, 1933 : 9), a fact taken into con-
sideration, with others, by McDowell & Bogert in aligning the Pygopodinae and
Lialinae with the Gekkota ; but it seems unlikely that development of these bones
to form external hooks would have occurred twice in Australasia, and nowhere else,
pari passu with reduction of limbs, the acquisition of a serpentiform habit and
sundry other features. So the position of Aprasia in the Pygopodidae seems
assured. Within this family it is unique in its reduced dentition, degenerate eye,
absent tympanum, rudimentary middle ear (Shute & Bellairs, 1953), the lack of a
splenial bone (McDowell & Bogert, fig. 21) and vestigial shoulder girdle ; the latter
is much more complete in both Pygopus and Lialis (Fiirbringer, 1870 : pi. i, figs.
5, 6 : Kinghorn, 1926). It differs from Lialis in its paired parietal bones and
resembles Pygopus, Paradelma, Delma and Pletholax in this respect. If, therefore,
subdivision of the Pygopodidae into subfamilies is desirable, there would be
justification for placing Aprasia in a distinct subfamily by itself. This procedure
would, however, tend to obscure the fact that the genera Pygopus, Paradelma,
Delma and Aprasia form an almost continuous linear series showing progressive
limb-reduction and shortening and consolidation of the skull. Aprasia is the most
completely " degenerate " of the series as it is also, probably, the most truly fossorial.
SPECIES, RACES AND VARIATION
As already mentioned, the material examined is divisible into five morphological
groups which form a rational geographical picture, briefly as follows :
I. A postocular shield present.
(a) Snout prominent (fig. 3A), 1-5 to 2-1 times as long as the eye. 1 A suture normally
runs from the nostril to the second labial, the nasal and second labial shields being
in contact.
(1) Presacral vertebrae 95-1 10 (mean 101-4). Mid -body scale rows 12 or 14.
Victoria, South Australia, Nurina (W.A.).
(1) Aprasia stviolata striolata.
(2) Presacral vertebrae 88-94 (mean 91-8). Mid-body scale rows 12. Albany,
Plantagent and Tambellup (W.A.).
(2) Aprasia striolata glauerti.
1 The method of measurement is important and is discussed below (p. 375).
37°
THE LIZARD GENUS APRASlA
(b) Snout less prominent (Text-figs. 3B and 6), i-i to i-8 times as long as the eye.
No suture running from the nostril. Presacral vertebrae 98-110 (mean 98-5) in
males and 102-105 (mean 103-5) in females. Mid-body scale rows 14. Albany to
Gerald ton (W.A.) : ? Northern Territory.
(3) Aprasia pulchella.
II. No postocular shield. Snout very prominent, 1-4 to 2-5 times as long as the eye (Text-
figs. 3c and 7). A suture from the nostril reaches the prefrontal so that the nasal and
second labial shields are not in contact or meet at a point only.
(a) Presacral vertebrae 96-112 (mean 102-8) in males and 98-116 (mean 108) in
females. Mid-body scale rows 12 (rarely 14). Albany to Shark Bay (W.A.).
(4) Aprasia repens repens.
(b) Presacral vertebrae 125, male and 137, female. Mid-body scale rows 14. Hermite
Island (W.A.).
(5) Aprasia repens rostrata.
Fig. 3. Three male specimens of Aprasia from Albany, to show lateral head scales
and proportions, a. Aprasia striolata glauerti subsp. nov. Holotype. b. Aprasia
pulchella Gray. Perth Mus. 10955. c - Aprasia repens repens (Fry). Perth Mus.
10953-
In this synopsis the assemblages have been given specific or subspecific status
and justification of these treatments is called for. It will be observed that many
of the characters used in the key " break down " if they are considered without
reference to geographical distribution. For instance, mid-bod}/ scale rows and
THE LIZARD GENUS APRASIA 371
numbers of presacral vertebrae show a total range of variation which permits of no
absolute differentiation between what are labelled as " species ", thus :
A . striolata.
A . pulchella.
A . repen.
Scale rows
12- 14
14
12- 14
Vertebrae
88-110
98-110
96-137
If, however, these characters are considered only in the geographical region where
the three are sympatric, i.e. the Plantagenet Division of Western Australia, they
provide in combination a clear cut differentiation, thus :
A . striolata. . A . pulchella. A . repens.
Scale rows 12 . 14 . 12
Vertebrae . . 88-94 • 98-103 . 96-106
A. pulchella is, additionally, completely differentiated from both of the others by
its much less prominent snout with its less extensive rostral (less than one-third
the diameter of the eye), the complete fusion of the nostril and first upper labial,
and a distinctive colour pattern. Additional differences between A. striolata
glauerti and A. repens repens exist in the same three features, viz. : Eye /rostral
ratio 1-5-1-9 in glauerti vice 1-2-1-7 in repens, the suture behind the nostril running
to the second labial in glauerti but to the prefrontal in repens and in colour. They
are also completely differentiated by the post-ocular condition. Thus, the Aprasias
of the Plantagenet Division can be subdivided into three different and distinct
morphological groups which are not associated with age, sex, season, etc. It is
reasonable to suppose that this fact signifies that there are three non-interbreeding
assemblages and the only logical taxonomic treatment is to regard them as three
distinct species. The most distinct morphologically is A. pulchella and this is
also the species with the smallest geographical range (from Albany to Geraldton)
and with the least variability. A . repens and A . striolata resemble each other more
than either does A. pulchella and both show geographical variation of the same
nature.
A. repens ranges northwards from Albany to the Monte Bello Islands and its
distribution along this range may prove to be continuous, though the samples
examined are not. The list of specimens examined, on pp. 382-383, is arranged in
approximately south-north order, and it will be observed that, when due allowance
is made for sexual and individual variation, there is a progressive increase in the
number of presacral vertebrae from south to north ; there is also an increase in
the number of mid-body scale rows from 12 to 14. It is, however, clear that the
variation is not exactly correlated with latitude. For instance, in a series of sixteen
males from Perth and its environs the vertebral range is from 95 to 108, whereas in a
series from around Northam in almost the same latitude the range is noticeably higher,
from 108 to 112. This and similar discrepancies may be due to chance and the
short series available, but it seems at least equally possible that some environmental,
possibly climatic, factor is involved. Although there is an overall increase in
temperature from south to north, the isotherms do not follow an east-west course ;
372
THE LIZARD GENUS APRASIA
Northam has a more " continental " climate than Perth with higher temperatures
in summer, when the isotherms are locally almost parallel with the coast (c.f. Year
Book of Australia, 40, 1954, p. 33). Rainfall, too, though generally decreasing with
decreasing latitude, also diminishes with distance from the coast. Unfortunately
climatological information is not available for many of the localities from which
specimens have been examined, and in any event the series are often very short.
But if localities are grouped into regions around climatological stations it is possible
to test, roughly, for correlation. The following table shows such groupings in
relation to the six stations named. No climatic data being available for the Monte
Bello Islands, the temperature and rainfall figures of Onslow have been used.
Rainfall
Temperature
Vertebrae.
(annual
(mean
.._,.„. ., A
average)
December)
r
■ '**
Station.
fr
?•
Scales.
(in).
(° F.).
Albany
96-0 (2)
—
12
39-7
64-2
Katanning .
97'3 (3)
104-4 (8)
12
19-4
68-3
Collie .
—
103-0 (2)
12
39-6
68-i
Perth .
102 -6 (20)
108-6 (25)
12
36-0
71*0
Geraldton .
107-0 (2)
II2-0 (i)
12/14 .
18-6
72-7
York .
110-7 (3)
iii-o (2)
12
18-0
73*5
Hermite Is.
125-0 (1)
I37-0 (I)
14
—
—
(Onslow)
—
—
9'4
83.0
here is no obvio
us correlation between num
bers of veri
:ebrae and
rainfall, either
annual or monthly. But Text-fig. 4, in which vertebrae are plotted against the
mean normal temperature for December, shows a correlation very clearly, though
what its significance may be remains to be discovered. It is, however, difficult
not to suspect that the correlation may be due to direct cause and effect, since it
has been shown experimentally that the numbers of meristic structures in poikilo-
thermic animals may be changed by varying the temperature during development.
Wade Fox (1948), for example, has shown that in a Garter Snake, the numbers of mid-
body scale rows, and ventral and subcaudal scutes are higher in groups of individuals
whose mothers were kept at higher temperatures during pregnancy. It has also
been established that in fishes (e.g. Taning, 1952, Lindsey, 1954) an increased
temperature during the appropriate embryonic period will, in some species, result in
an increased number of vertebrae, fin rays, etc. (and vice versa) within certain
limits ; a decrease in temperature below a certain point may also, sometimes,
result in an increase of the meristic characters. Detailed information regarding the
breeding season of Aprasia in all areas, together with temperature records of the egg
sites will be necessary to establish that we are, in fact, dealing here with a thermo-
plastic effect. In the meantime all that can be said is that there is evidence of a
cline correlated with the temperature in summer, a time of year when breeding
may be expected to occur. There is a discontinuity in the series and it is, therefore,
legitimate to subdivide it at the discontinuity, treating the two parts as subspecies.
A. striolata, as already mentioned, also shows geographical variation in vertebral
and mid-body scale numbers. It ranges from the south of Western Australia,
THE LIZARD GENUS APRASIA
373
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374
THE LIZARD GENUS APRASIA
where it is sympatric with A. repens in Albany, Plantagenet and Tambellup, across
the south of the continent to western Victoria. In the west its vertebrae range
from 88 to 94, there being no significant secondary sex difference, and the mid-body
scales are constantly 12. In South Australia and Victoria the vertebrae range
from 95 to no and 25% of the specimens seen have 14 scale rows. Again, as in
A. repens, there appears to be some correlation with temperature, but not with
rainfall, thus :
Rainfall
Temperature
(annual
(mean,
average)
December)
Station.
Vertebrae.
Scales.
(in.).
(° F.).
Albany .
92-0 (11)
12
39'7
64-2
Katanning
9i-3 (3)
12
19-4
68-3
Eyre (Nurina)
IOI'O
14
n-6
67.0
Wudinna 1
99-0
14
15-0
71.0
Adelaide
101 -4 (10)
12 or 14
21 -o
70-7
1 Climatic data estimated as intermediate between Port Lincoln and Port Augusta.
There is, however, inadequate information to establish the existence of tempera-
ture plasticity and, moreover, a marked discontinuity becomes obvious when the
series is plotted (Text-fig. 5). So in the present state of knowledge the recognition
of two subspecies is indicated.
75
70
65
• Eyres Penins.
" Adelaide
° O
- Katanning
O
- Eyre (Nurina)
O
- Albany
O
Subsp. glauertl
1 1 1
Subsp. itrlolata
I > 1 1
90
92
94
96
Vertebrae
98
100
102
Fig. 5. Aprasia striolata. Average numbers of presacral vertebrae of local populations
plotted against the mean December temperatures of the areas,
THE LIZARD GENUS APRASIA 375
Only sixteen specimens of A . pulchella with definite localities have been examined,
all from points between Albany and Geraldton. Within this restricted area there
appears to be little variation ; all the specimens have 14 scale-rows and the presacral
vertebrae range between 96 and 98 in males (8) and from 102 to 105 in females (8).
The type, a female without precise locality, has 104. The species may have a greater
geographical range and there may be some geographical or temperature variation ;
one specimen seen (Sydney, No. 12529) is an immature male with a much higher
number of vertebrae, no. This specimen has no locality recorded against it in
the registers of the Australian Museum, but formed part of a mixed collection
mainly from the Northern Territory (Yirrikala Mission) and the northern parts of
Western Australia.
The number of scales longitudinally is associated with the number of vertebrae
and consequently shows the same geographical variation within each species. There
are, however, differences between the species in regard to the number of scales in
relation to the number of vertebrae, i.e. relative size of the scales, and the differences
appear to be significant. The ratio of the number of ventral scales to the number
of vertebrae varies as follows :
A. pulchella (18) 1*44 to 1-59. Mean 1-52 00-049.
A.striolata (28) 1 -31 to 1 -50. ,, 1-4300-048.
A.repens (48)1-23101-52. ,, 1-3500-06.
Comparing the three on the criterion of the quotient of the difference of the means
divided by the standard error of the difference of the means, the differences are all
significant, thus :
(a) A. pulchella j A. striolata d = 6-4.
~Vd
(b) A. striolata j A.repens ,, = 6-5.
(c) A. pulchella I A.repens ,, = 11 «6.
A. striolata is almost exactly intermediate between the other two, A. pulchella
having the smallest scales. Increase in scale-size is often accompanied by complete
or partial elimination of some scales. In Aprasia this effect is manifest in the head
region. In A. repens, with the largest scales, the post-ocular has been eliminated
and the pre-ocular is much reduced in size compared with the other two species
(c.f. Text-figs. 3, 6 and 7). No variation in this character has been observed.
The character of length of snout relative to size of eye can be appreciated by
inspection, but is difficult to measure accurately. The method adopted was to
measure the distance between the tip of the rostral and the anterior corner of the
frontal as seen in plan projection on a plane parallel with that passing through
the tip of the rostral and the centres of the eyes, using a camera lucida for the
purpose. The eye diameter is that of the brill measured horizontally, i.e. excluding
the ring of small scales that borders the " window ". The method cannot be claimed
to give a high degree of accuracy, but it does provide directly comparable figures
376 ^THE LIZARD GENUS APRASIA
although the range of variation will almost certainly be exaggerated. Comparison
between the subspecies shows no significant differences, thus :
(a) A. striolata striolata (n). Mean i -67 o 0-14^! d = 1 -62.
A. striolata glauerti (14). ,, 1-78 oo-i6j ^
(b) A. repens repens (65). ,, 1-96 o o-2o^\d = < 1,
A . repens rostrata (2) . „ 2-1 ? J ad
Between species, however, the differences are marked and significant, thus :
A.pulchella (17). Mean 1-45 00-15.
A. striolata (25). ,, 1-74,, 0-16.
A. repens (67). ,, 1-96, ,0-20.
(a) A. pulchellajA. striolata d = 6-o.
ad
(b) A. striolata I A. repens ,, = 5-5.
(c) A. pulchella/A. repens ,, = 11-9.
The character of the length of the rostral shield, as might be expected, is correlated
with the length of the snout ; it shows no infraspecific variation that is certainly
significant. As between species, however, the ratio of brill diameter to length of
rostral* is :
A. pulchella (17). Mean 3-90 o 0-53.
A. striolata (25). „ i- 80 00-14.
A. repens (65). ,, 1-5200-18.
Comparing the differences of the means for significance :
(a) A . pulchella I A . striolata d =16-2.
ad
(b) A. striolata J A. repens ,, = 8-8.
(c) A. pulchella /A. repens ,, = 19-0.
Thus, it seems to be established that infra-specific variation occurs mainly, if
not exclusively, in meristic characters — the numbers of vertebrae and scales. Non-
meristic characters such as the length of the snout relative to the eye (and the asso-
ciated character of the relative proportions of the rostral shield), the presence or
absence of a post-ocular shield, and relative scale size are significantly different
between the samples that are believed to represent bio-species, but show no signi-
ficant geographical variation within these groups.
SYNONYMIES AND DESCRIPTIONS
Aprasia striolata striolata Liitken
Aprasia pulchella var. striolata Liitken, 1863, Vidensk. Medd. Kbh. 1862 : 300, pis. 1-2, figs. 3-3C
Aprasia pulchella var. lineolata Liitken, 1863, loc. cit.
Aprasia octolineata Peters, 1864, Monatsber. Ak. Berlin, 1863 : 233.
Aprasia pulchella, Giinther, 1873 Ann. Mag. nut. Hist. (4) 12 : 145 (part). Giinther, 1875, Zool.
* (Measured in plan projection.)
THE LIZARD GENUS APRASIA
377
Erebus and Terror, 2 : 10 (part). Boulenger, 1885, Cat. Lizards Brit. Mus. 1 : 246 (part).
McCoy, 1888, Prodrom. Zool. Victoria, 17 : 233, pi. 161, fig. 1. Lucas & Frost, 1894, Proc. roy.
Soc. Vict., n.s., 5 : 340. Lucas & le Souef, 1909, The Animals of Australia (Melbourne) : 219
(part). Werner, 1912, Das Tierreich, 33 : 25 (part), fig. 5. Zietz, 1920, Rec. S. Aust. Mus.
1 : 193 (part). Kinghorn, 1923, Rec. Aust. Mus. 14, 2 : 130 (part), figs. 7-9. Kinghorn,
1926, Rec. Aust. Mus. 15 : 63, fig. 18. Kershaw, 1927, Vict. Nat. 43 : 341. Waite, 1929,
Rept. and Amph. of South Australia (Adelaide) : 95, figs. 69, 70
Ophiopsiseps nasutus (non Bocage), Jensen, 1901, Vidensk. Medd. Kbh. 1900 : 317, pi. 3.
Ophioseps nasutus, {non Bocage), Fry, 1914, Rec. W . Aust. Mus. 1 : 181 (part). Kinghorn, 1923,
Rec. Aust. Mus. 14, 2 : 128 (part), figs. 4-6. Kinghorn, 1926, op. cit., 15 : 62, fig. 17.
Snout projecting and rounded, its length 1-5 to 2-0 times as long as the horizontal
diameter of the brill (mean 1-67, cr 0-14). Rostral well visible from above, its
length contained i-8 to 2-0 times in the length of the brill (mean 1*90, cr 0-07). A
suture normally connects the nostril with the suture between the first and second
labials so that the nasal and the second labial are in contact ; in one specimen
only of the fifteen examined are these two shields separated. A small postocular
separates the fourth labial from the supraocular. Premaxillary teeth may be
present, though small, in females ; they are constantly present, and much larger,
in males. Scales in 12 (80%) or 14 (20%) longitudinal rows at mid-body ; from
140 to 155 scales, approximately, in a straight line from the mental shield to the
vent in the proportion of 1*38 to 1-50 (mean 1*43, cr — 0-034, n. 13) per vertebra.
Presacral vertebrae number 98 to 104 in males (5 counted) and 95 to no in the nine
females. A well defined colour pattern of dark lines or rows of dots is usually
present on the back. The largest female examined measures 133 mm. from snout
to vent and the largest male 107 mm.
This subspecies appears to range from western Victoria westwards to the Nurina
Division of Western Australia. The reasons for its recognition have been discussed
above. The material examined includes the immature specimens which Jensen
misidentified as Ophioseps nasutus Bocage. The variation in the more important
taxonomic characters is as follows :
Verte- Snout. Eye
Museum No.
Locality.
Cop. R.50.2
Hochkirk, Vict.
Sydney 11 759 •
Jeparit, Vict.
11760 .
,, ,,
B.M. 1902.7.30.8
Renmark, S.A.
,, 1902.5.30. 10
>> >>
,, 1902.5.30. 11
y, >,
,, 1902.5.30. 12
,, ,,
„ 1902.5.30. 13
,, >,
,, 1862.7. 19.4
S. Australia
,, 1862.7. 19.5
,, ,,
,, 1862.7. 19.6
,, ,,
Sydney 14382 .
Wudinna, S.A.
3466 .
p
B.M. 46.5.2.14
?
Perth R.5280 .
Eyre, Nurina, W.A
Sex.
Length.
Scales.
brae.
?
57
12
100
• ?
?
12
?
• $
?
12
95 •
• ?
79
12
102
■ $
no
14
109
• ?
109
14
IOI
• 6*
100
12
98 .
■ 6*
95
12
IOI
■ 0*
102
12
104
• ?
118.
12
104
• 6*
no
12
IOO
■ <J
107
14
99 •
• ?
115
12
102
• $
133
12
no
• ?
113
14
IOI
Eye
i-7
?
?
i-5
1-7
2'0
i-5
1-7
i-6
1-6
?
?
i-7
i-6
i-8
Rostral.
i-8
378
THE LIZARD GENUS APRASIA
Aprasia striolata glauerti subsp. nov.
(Text-fig. 3A)
Snout projecting and rounded (Text-fig. 3A), its length i-6 to 2-1 times as long
as the horizontal diameter of the brill (mean 178, or = 0-16). The rostral is well
visible from above, its length being contained 1-5 to 1-9 times in the length of the
brill, the mean being 1-70 and <r 0-14. The condition of the nasal suture is similar
to that of the eastern race, but two specimens of the fifteen are asymmetrical and
have the nasal separated from the second labial on one side ; in one specimen the
suture is incomplete and fails to reach either the interlabial or the naso-prefrontal
suture. Mid-body scale rows are constantly 12 and there are from about 120 to 140
from the mental shield to the vent in the proportion of 1-31 to 1-49 (mean 1-43, a* 0-058,
n. 15) per vertebra. Presacral vertebrae number 88 to 93 in males (7) and 91 to
94 in females (8). Premaxillary teeth present in both sexes, but smaller and some-
times absent in females. Straw coloured above and beneath, sometimes with
obscure darker markings suggesting dusky lines on the back. The largest male
measures 109 mm. from snout to vent and the largest female 115 mm.
The type series is from the " road districts " of Albany, Plantagenet and Tam-
bellup, as listed below, the first mentioned being the holotype.
Museum No. Locality.
Sex.
Length.
Vertebrae.
Perth 10949
Albany
6*
79
93
„ 6782
>»
9
89
92
„ 6782
>»
6*
89
88
„ 6782
>>
$
89
9i
„ 6782
,,
?
95
94
,, 6782
' »»
6*
97
93
„ 6782
y,
?
98
93
„ 10950
,,
9
115
92
„ 1 095 1
>,
9
92
9i
B.M. 1911.12.
18.3.
9
108
93
Perth 7214
Narrikup
6*
84
93
Sydney 12305
Tambellup
0"
79
90
12305
•
«J
> 88
92
12305
,,
$
92
92
n 3468
. [" Australia "]
6*
109
92
Apra
(Te
sia pw
xt-figs.
Ichella Gra
3B and 6)
y
Snout
Eye
Eye
Rostral.
I
9
1 •
I
7
1 •
I
6
1 •
I
9
T •
I
9
I •
I
8
I •
I
7
I *
2
1
I •
I
9
I •
I
7
I •
2
I •
I
6
I '
I
6
I •
I
6
I •
)
?
Aprasia pulchella Gray, 1839, Ann. Nat. Hist. 2 ; 332. Gray, 1841, in Grey, Travels in Australia,
2 : 428, 438, pi. 4, fig. 2. Gray, 1845, Cat. Lizards Brit. Mus. : 68 (part). Giinther, 1873,
Ann. Mag. nat. Hist. (4) 12 : 145 (part). Giinther, 1875, Zool. Erebus & Terror, 2 : 10 (part).
Boulenger, 1885, Cat. Lizards Brit. Mus. 1 : 246 (part). Lucas & le Souef, 1909, The Animals
of Australia, (Melbourne) : 219 (part). Zietz, 1920, Rec. S. Aust. Mus. 1 : 193 (part).
Kinghorn, 1923, Rec. Aust. Mus. 14, 2 : 130 (part). Kinghorn, 1926, Rec. Aust. Mus. 15 : 63
(part). Loveridge, 1934, Bull. Mus. comp. Zool. Harv. 77 : 316.
THE LIZARD GEiNUS APRASIA
379
Ophioseps nasutus Bocage, 1873, /. Set. math. phys. nat. Lisboa, 4 : 232. Werner, 191 2, Das
Tierreich : 33: 25. Fry, 1914, Rec. W. Aust. Mus. 1 : 181 (part). Kinghorn, 1923, Rec. Aust.
Mus. 14, 2, 128 (part).
Ophiopsiseps nasutus, Boulenger, 1887, Cat. Lizards Brit. Mus. 3 : 436.
Aprasia brevirostris Werner, 1909, in Michaelsen & Hartmeyer, Die Fauna sudwest- Austral.
2, 16 : 266, fig. 2.
Snout slightly prominent and bluntly rounded (Text-figs. 3B and 6), its length
i-i to i-8 times the length of the brill (mean 1-45, a 0-15). The portion of the
rostral visible from above is less extensive than in any other species, its length
being contained 3-1 to 5-0 times in the brillar length (mean 3-90, cr 0-53). No
suture between the nasal and first labial shield, though in the holotype there is a
vestige on the nasoprefrontal suture (Text-fig. 6). A postocular separates the
fourth upper labial from the supraocular. Mid-bod}/ scale rows 14 ; from 145
to 170 scales, approximately, on the mid-line between the mental shield and the
vent in the proportion of 1-44 to 1-59 (mean 1-52, cr 0-049 > n - z &) P er vertebra.
Presacral vertebrae 96 to no in males (9) and 102 to 105 in females (9) ; the possi-
bility of geographical variation in this character has already been discussed. Pre-
maxillary teeth absent in females. Pale brown above with indefinite dusky linear
markings on the dorsal scales which can produce a lineolate appearance ; on the
head the dark markings are more irregular and produce a vermiculate appearance.
The largest male examined measures 108 mm. from snout to vent and the largest
female 124 mm.
This species is known with certainty to occur from Albany to Geraldton.
Although Boulenger (1885) lists two specimens as " types " of A. pulchella, there
is nothing in Gray's original description to indicate that he had more than one
specimen whose dimensions are given as 2f in. from snout to vent with a tail of
if in. Only one of Boulenger's specimens, the one listed below as B.M. 1946 . 8 . 30 . 93,
Fig. 6. Aprasia pulchella Gray. Holotype female.
3 8o
THE LIZARD GENUS APRASlA
approximates to these dimensions (snout to vent 2-8 in., tail 175 in.) and has ver-
miculate head-markings (c.f. Text-fig. 6) as figured by Gray in 1841. The only
doubts on the matter arise from the facts that (1) the original description states that
the nostril lies " in the suture between the top of the first labial and anterior frontal
plate " and (2) Gunther in 1873 stated that all the specimens then in the British
Museum had only 12 mid-body scales. In the specimen now regarded as the type
the first labial is fused with the shield above it and there are 14 scale rows. Gunther
was patently in error, as was Boulenger (1885), also, when he failed to detect more
than 12 scale rows and it seems likely that Gray, using only a hand lens, was misled
by the vestige of a suture.
Thanks to the courtesy of Professor Jorge the holotype of Ophioseps nasutus
Bocage has been examined and compared with the other specimens listed. Unfor-
tunately the specimen is in very poor condition ; it is in four fragments, breaks
occurring after the ninth vertebra, in front of the nineteenth presacral, and imme-
diately behind the pelvic girdle ; the tail tip is also wanting and the scales of the
upper surface of the head are very badly abraded, so that their limits can only be
determined with difficulty and a little uncertainty. It is also likely that the specimen
has been partially desiccated at some time, since the cranium was full of air and
the whole specimen measures only 73% of its pristine length (head and body frag-
ments 95 vice 133 mm. and tail 38 vice 52 mm.). 1 Nevertheless, despite these
defects, a sufficient number of characters can be deciphered or measured to make it
virtually certain that the species involved is the same one which Gray had previously
named Aprasia pulchella. Thus, for comparison with the characters of the species :
0. nasutus
Aprasia
Suture from nostril .
(Type).
Absent
pulchella.
Absent
Snout length /brill diam.
Brill diam. /rostral length
Postocular
1-42
3'7
Present
Mean 1-45
„ 3'9
Present
Mid-body scales
Scales/vertebrae
14
i'57
14
Mean 1 -52
As noted above, the skull is visible dorsally and the ratio of frontal to nasal length
(1*56) is similar to that in an example of A. pulchella (1*50). The presacral vertebrae
number 89, but it is certain that a number have been lost from the ends of most
of the fragments (one was found loose) and no significance can be attached to the
number. The sex cannot be determined since, although no post-cloacal bones are
present, there is a clear indication of the loss of at least 4 anterior caudal vertebrae.
The first of the tail vertebrae has a well-defined fracture-plane through its centrum
and so cannot be one of the most anterior caudal vertebrae which possess $0
such split (Pratt : 184) ; in the numerous X-ray photographs of Aprasia that
have been examined the fracture-plane is not evident anterior to the fifth to
seventh caudal.
1 The reduction cannot be due to the loss of a single large fragment of 52 mm. because when the tail
and the head-plus-body are considered separately each is shorter than originally in the same proportion.
THE LIZARD GENUS APRASlA
38i
The specimens examined and their characters are :
Verte- Snout.
Eye
Museum No.
Locality.
Perth 10954 • • • Albany
10955
,,
» 2371
Mt. Baker
10711
Boddington
10712
,,
642
Wagin
„ 3873
Lowden
1984
Mt. Helena
5994
Statham
4629
Huntley
» 3342
Darlington
4629 A
Guildford
M.C.Z. 24460
Mundaring Weir
Perth 3409 .
Upper Swan River
B.M. — .
Swan River
M.C.Z. 24467
Geraldton
24468
,,
B.M. 1946.8.30.9
3
W. Australia
Sydney 6553
?
12529
?
Mus. Bocage
" Australia "
Sex.
Length.
brae.
Eye
$
117
102
1-6 .
• 0"
92
98
1-6 .
• $
109
103
i-4 .
• S
95
96
i'5 •
• o*
108
97
1-6 .
• 9
114
104
1-4 .
. 6*
74
96
i'5 •
• $
p
104
i-8 .
• 0*
99
98
i-5 •
• 0*
86
97
i-4 .
• $
106
105
i-4 .
• $
104
103
i-4 .
• 6*
—
98
i'3 •
• ?
124
105
i'5 •
• $
106
?
1 -i
• $
—
103
1-6 .
• o*-
—
98
i-5 •
• ?
72
104
I -2
?
115
?
. i- 4 .
• S
62
no
1-4 .
?
95 +
89 +
1-42 .
Rostral.
3*2
3'8
3*1
5-o
4-0
4'2
3-3
3-6
4-0
4'5
4-0
3'5
3-S
3-6
3-5
4-7
4'5
3'7
Aprasia repens repens (Fry)
(Text-fig. 3c)
Aprasia pulchella (part), Gray, 1845, Cat. Lizards Brit. Mus. : 68. Giinther, 1875, Ann. Mag.
nat. Hist. (4) 12 : 145. Giinther, 1875, Zool. Erebus & Terror, 2 : 10. Boulenger, 1885, Cat.
Lizards Brit. Mus. 1 : 246. Werner, 1909, in Michaelsen & Hartmeyer; Die Fauna siidwest
Austral. 2, 16 : 266. Lucas & le Souef, 1909, The Animals of Australia (Melbourne) : 219.
Werner, 1912, Das Tierreich, 33 : 25. Zietz, 1920, Rec. S. Aust. Mus. 1 : 193.
Ophioseps repens Fry, 191 2, Rec. W. Aust. Mus. 1 : 178, fig.
Aprasia repens, Kinghorn, 1923, Rec. Aust. Mus. 14, 2 : 132, figs. 10-12. Kinghorn, 1926,
Rec. Aust. Mus. 15 : 64. Loveridge, 1934, Bull. Mus. comp. Zool. Harv. 77 : 317.
Lialis repens, Kinghorn, 1926. torn. cit. fig. 19.
Snout strongly projecting (Text-fig. 3c), its length 1-4 to 2-5 times as long as the
brill (mean 1-96, cr 0-20). Rostral well visible from above, its length contained
i-o to 2-4 times in the brillar diameter (mean 1-52, a 0-18). A suture connects the
nostril with the anterior border of the prefrontal so that the first labial and the
prefrontal are in contact ; in two specimens the four shields (nasal, prefrontal,
first labial, second labial) meet at a point and in another example the two sides arc
not symmetrical the nasal being in contact with the second labial on the right
side but separated from it on the left. There is no post ocular, the fourth labial
reaching the supraocular. Premaxillary teeth absent in females. Scales in 12
(75 specimens) or 14 (1 specimen) rows at mid-body and between 125 and 165,
approximately in a median line from the mental shield to the vent in the proportion
of 1-23 to 1-52 (mean 1-35 ; cr 0-06 ; n. 46) per vertebra. Presacral vertebrae
3«2
THE LIZARD GENUS APRASIA
96-112 in males and 98-116 in females, there being geographical, possibly temperature
correlated, variation as discussed above. The ground colour is pale straw, above
and below, often with ill-defined dark longitudinal lines or rows of dots on the back
and especially on the flanks ; head sometimes with dark markings ; upper lip and
throat chrome yellow ; tail red or yellow. The largest male examined is 150 mm.
from snout to vent and the largest female 165 mm.
The subspecies ranges from Albany to Victoria (W.A.) and possibly further
north. Werner (1909) records a specimen in the Perth Museum as from Shark Bay,
but there is no present record of such a specimen. The only possibility is specimen
R.i 1360, a male which bears a label " M & H " (= Michaelsen and Hartmeyer) ;
but Werner's record is not listed as having been obtained at any of the stations
where Michaelsen and Hartmeyer worked and which are all numbered. In addition
its vertebral count (104) is much more compatible with an origin near Perth.
The material examined contained these specimens :
Verte-
Snout.
Museum No.
Locality.
Sex.
Length.
brae.
Scales.
Eye
Perth 10952 . . . Albany
6*
73
96
12
18
„ io953
"
6*
98
96
12
2-1
B.M. 1931.7.1.139
Mt. Toolbrunup
6*
95
96
12
i-7
Perth 2868
Tambellup
$
104
104
12
i"7
„ 2869
,,
83
99
12
19
„ 2870
,,
$
93
103
12
1-9
„ 2871
,,
$
95
103
12
1-8
2872
,,
$
95
104
12
1-8
10692
Borden
?
116
106
12
1-8
.. 433^
Dumbleyung
0*
82
97
12
2-1
.. 433B
„
9
103
105
12
2-1
» 373 •
,,
$
105
106
12
2'3
Sydney 8015
y »
$
IOI
104
12
?
Perth 10246 . . . Williams
?
107
108
12
2-0
10322 . . . Bunbury
?
95
98
12
1-9
10895 . . West Midland
6*
82
IOI
12
2-1
10894
,, ,y
$
90
107
12
1-9
„ 10893
y> »>
?
82
in
12
i-8
» 4173
Midland
?
93
107
12
2-1
„ 766 .
Midland Junction
?
107
no
12
2-0
„ 3734
Rottnest Isld.
-A
O
62
107
12
2-3
„ 3759
>, 11
O
?
92
12
2-1
» 3765
.» y,
?
63
III
12
1-9
» 4582
y, ,y
?
109
113
12
2-3
M.C.Z. 33027
,,
O"
—
no
12
—
33° 28
,,
$
—
113
12
—
33029
,,
?
—
109
12
—
33031
,,
$
—
no
12
—
33032
,,
? •
—
no
12
—
THE LIZARD GENUS APRASIA
383
Museum No.
Perth 4373
B.M. 69.5.25.18
Perth 3347
„ 2596
.. 1742
,, 6944
„ 631
2251
Perth 9664
„ 9665
„ 2837
.. 8745
„ 8185
„ 9093
„ 9094
>> 5344
M 123 •
» 7405
„ 74O6
M 7390
„ 53IO
„ 8721
I0208
2I50*
» 1374
„ 5351
» 2397
„ 4186
» 707 •
» 4383
M.C.Z. 24427
Perth 4356
„ 4357
Sydney 9138
Perth 5492
„ 8891
» 4913
„ 6927
9222
„ 1730
„ 5064
M.C.Z. 24458
24459
B.M. 43.5.19.86
Perth 1 1360
Locality.
East Perth
Perth
Rivervale
Gooseberry Hill
Claremont
Nedlands
Cottesloe
Applecross
Ba}'swater
Bassendean
Leederville
Maylands
Mt. Lawley
E. Freemantle
Balcatta Beach
Stoneville
Mundaring
Northam
Bakers Hill
Lomos
Newmarracarra (Vict.
Eradu (Vict.)
Geraldton
W. Australia
?
Sex.
6*
6*
S
s
?
?
6*
6*
$
$
6*
6*
?
o*
(J
?"
6*
o*
?
?
?
?
?
cJ
6*
?
?
o*
?
?
$
?
6*
o*
6*
?
Length.
107
100
96
87
104
76
52
98
88
55
87
no
78
80
7i
58
105
80
72
68
92
98
102
103
102
7i
82
57
47
106
102
63
75
123
92
102
93
80
?
60
Verte-
brae.
105
?
105
101
106
103
?
105
IOI
103
114
114
IOO
102
102
104
106
95
104
99
108
108
108
107
113
109
106
101
102
106
105
108
?
106
116
112
112
108
106
?
109
Scales.
— • 105
85
60
?
104
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
12
14
12
12
12
12
Snout.
Eye
2-0
2-3
2-0
1-9
2-1
2-0
2-0
19
2-3
I
7
o
o
7
o
I
2
8
2-1
2 -I
1-8
2 -O
2.0
2-0
2-0
1-7
1*5
1.4
1-7
1-9
i-6
2-1
2-0
1-6
1-9
2 -o
2-0
2-2
2-5
2-0
2-0
* This specimen is ovigerous.
3«4
THE LIZARD GENUS APRASIA
Aprasia repens rostrata subsp. nov.
(Text-fig. 7)
Aprasia sp. Hill, 1955, Proc. Linn., Lond. 165, 2 : 115.
Holotype a male, collected 17.viii.52 on Hermite Island and allotype a female
collected on the same island 22. v. 52.
Snout very strongly projecting (Text-fig. 7) its length 2-0 times and 2-2 times as
long as the diameter of the brill in male and female respectively. Rostral shield
drawn out posteriorly so that the length of the portion visible from above is contained
0-9 (<?) and o-8 (?) times in the brillar length. The suture running backwards from
the nostril meets the suture between the first and second labials in such a position
Fig. 7. Aprasia repens rostrata subsp. nov. Allotype female.
that these two labials, the nasal and the prefrontal meet at a point. No post-
ocular shield, the supraocular produced ventrally to make contact with the fourth
labial. Premaxilliary teeth present in the male but absent in the female. Scales
in 14 longitudinal rows at mid-body. Scales on the mid-line between the mental
shield and the vent are about 175 in the male and 190 in the female, counted to the
nearest five. Presacral vertebrae 125 in the male and 137 in the female.
Pale brown above and white beneath. A broad darker lateral band on the sides
of the neck which breaks up posteriorly into three fine brown lines. Three additional
fine brown lines on the nape, but the mid-dorsal region of the body not lined ; a
single mid-dorsal dark line along the tail. Another brown line on each side of the
belly. Head vermiculate.
Dimensions :
cJ. Snout to vent 99 mm. ; tail 65 mm. (incomplete).
?. „ >, » 109 mm. ; „ 13 mm. ( „ ).
THE LIZARD GENUS APRASIA 385
ACKNOWLEDGMENTS
Thanks are due to Sir William Penney who, despite his many preoccupations
with the major objects of the expedition to the Monte Bello Islands, arranged for
collections of the fauna and flora of the islands to be made, and to Surgeon
Commander G. Wedd, R.N., who organized and supervised these activities. The
resulting material touched off the present investigation, but no progress could have
been made without the assistance of Mr. L. Glauert, Mr. J. R. Kinghorn, Mr. F. W.
Braestrup and Mr. A. Loveridge, who have been most generous in lending comparative
material from their museums at Perth, Sydney, Copenhagen and Harvard. The
author is also grateful to his colleagues Mr. N. B. Marshall, and Mr. D. W.
Tucker, for help with ichthyological references, and to Mr. P. E. Purves for the
large number of X-ray photographs that had to be taken.
REFERENCES
Boulenger, G. A. 1 885. Catalogue of Lizards in the British Museum {Natural History).
Second edition.
Fox, Wade. 1948. Effect of Temperature on Development of Scutellation in the Garter
Snake, Thamnophis elegans atratus. Copeia, 1948 : 252-266.
Furbringer, M. 1870. Die Knochen unci Muskeln der Extremitdten bei den Schlangendhnlichen
Sauriern. Leipzig, 136 pp., 23 pis.
Gray, J. E. 1841. A Catalogue of the Species of Reptiles and Amphibia, hitherto described
as inhabiting Australia, with a description of some New Species from Western Australia
and some remarks on their geographical distribution. Appendix E. to Grey, G., Journals
of two Expeditions' of Discovery in North-West and Western Australia, 2 : 422-449.
Gunther, A. 1873. Notes on, and descriptions of, some Lizards with Rudimentary Limbs
in the British Museum. Ann. Mag. nat. Hist. (4), 12 : 145-148.
Jensen, A. S. 1901. Om Ophiopsiseps nasutus du Bocage, etc. Vidensk. Medd. naturh.
Foren. Kbh. 1900 : 317-328, pi. 3.
Kinghorn, J. R. 1923. Studies in Australian Reptiles, No. 3 : On the genera and species
of Ophioseps and Aprasia. Rec. Aust. Mus. 14, 2 : 126-134.
1926. A brief review of the family Pygopodidae. Rec. Aust. Mus. 15 : 40-64.
Lindsey, C. C. 1954. Temperature controlled meristic variation in the Paradise Fish
Macropodus opercularis (L.). Canad. J. Zool. 32 : 87-98.
Loveridge, A. 1934. Australian Reptiles in the Museum of Comparative Zoology, Cam-
bridge, Mass. Bull. Mus. comp. Zool. Harv. 77 : 316-317.
Lutken, C. 1863. Nogle nye Krybdyr og Padder. Vidensk. Medd. naturh. Foren. Kbh.
1862 : 292-311, pis. 1-2.
McDowell, S. B. and Bogert, C. M. 1954. The Systematic Position of Lanthanotus and
the Affinities of the Anguinomorphan Lizards. Bull. Amer. Mus. nat. Hist. 105 : 1.
Pratt, C. W. M. 1946. The plane of fracture of the caudal vertebrae of certain Lacertilians.
/. Anat., Fond. 30 : 184-188.
Shute, C. C. D. and Bellairs, A. d'A. 1953. The cochlear apparatus of Geckonidae and
Pygopodidae and its bearing on the affinities of these groups of lizards. Proc. Zool. Soc.
Lond. 123 : 695-709, 2 pis.
Smith, M. A. 1933. Remarks on some Old World Geckoes. Rec. Indian Mus. 35 : 9-19.
Vedel Taning, A. 1952. Experimental study of meristic characters of fishes. Biol. Rev.
27 : 169-193.
Werner, F. 1909. Reptilia exkl. Geckonidae und Scincidae. Fauna Stidw est- Aust. 2, 16,
251-278, pis. 14-15.
191 2. Eublepharidae, Uroplatidae, Pygopodidae. Fierreich, 33 : ix + 33.
PRINTED IN GREAT BRITAIN BY
ADLARD AND SON, LIMITED
BARTHOLOMEW PRESS, DORKING
S APR 1956
BIRDS COLLECTED BY
MR. F. SHAW-MAYER
IN THE CENTRAL HIGHLANDS OF
NEW GUINEA
1950-1951
R. w. SIMS
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 3 No. 10
LONDON : 1956
BIRDS COLLECTED BY MR. F. SHAW-MAYER
IN THE CENTAL HIGHLANDS OF NEW GUINEA
1950-1951
BY
R. W. SIMS
Pp. 387-438 ; Plates 13-14 ; 2 Text-figures
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 3 No. 10
LONDON: 1956
THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, is
issued in five series corresponding to the Departments
of the Museum, and an Historical Series.
Parts will appear at irregular intervals as they become
ready. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.
This paper is Vol. 3, No. 10 of the Zoological series.
PRINTED BY ORDER OF THE TRUSTEES OF
THE BRITISH MUSEUM
Issued March, 1956 Price Fifteen Shillings
BIRDS COLLECTED BY MR. F. SHAW-MAYER
IN THE CENTRAL HIGHLANDS OF NEW GUINEA
1950-1951
By R. W. SIMS
(with Field Notes and Photographs by W. T. Loke)
(Received 13th July, 1955)
CONTENTS
Page
Introduction ........... 389
Collecting Localities ......... 391
Annotated List of Birds Collected in the Wahgi Region by Mr. F.
Shaw-mayer .......... 393
References ........... 437
SYNOPSIS
This is a report on the birds collected by Mr. F. Shaw-Mayer in the Central Highlands of
New Guinea, mainly in 1950 and 1951. Two-hundred and seventy- two specimens were
obtained in the Wahgi region, principally on Mount Giluwe. Eighty-eight species and races
are represented of which one race was new to science.
INTRODUCTION
i. General
Mr. F. Shaw-Mayer has lived and collected for many years in New Guinea and has
travelled widely in the eastern half of the country. In recent years he has travelled
mainly in the Central Highlands and a collection of birds from that area forms the
subject of this report.
The Central Highlands of New Guinea were almost inaccessible prior to 1945 and
little was known of the region, its inhabitants or its fauna. In recent years, however,
the establishment of an airfield at Nondugl has made it more accessible and advantage
has been taken of this for making collections by others besides Mr. Shaw-Mayer. In
1950, and again in 1952, Mr. E. T. Gilliard of the American Museum of Natural
History made large collections of birds in the Wahgi region ; and during the inter-
vening year, 195 1, Count Gyldenstolpe also visited the area. The reports on these
ZOOL. 3, IO. 25
39°
BIRDS COLLECTED BY MR. F. SHAW-MAYER
expeditions were in the course of preparation when I started identifying and making
notes on Mr. Shaw-Mayer's specimens. The appearance of the first (Mayr and Gilliard,
1954) greatly facilitated the completion of these notes. The facts recorded below and
the taxonomic conclusions arrived at differ for the most part only in points of detail,
such as ranges of dimensions and various minor aspects of plumage colour, differences
which one would expect to find in two sets of random samples from the same area.
There are, however, a number of additional facts some of which suggest slight
modification to Mayr and Gilliard's conclusions. These notes had been completed
when I learnt that Count Gyldenstolpe's report was in the press. When it was
published (Gyldenstolpe, 1955) I read it in conjunction with my own notes and
referred to many specimens again. Generally there was a large measure of agreement
but on one or two points our conclusions were not identical, mainly because of
differences in age or sex of the specimens, or as a result of making comparisons with
material from different areas.
Mr. Shaw-Mayer gave brief field notes on some specimens and a number of these
are quoted here. Additional important field notes were made by Mr. Loke Wan Tho of
Singapore. Mr. Loke joined Mr. Shaw-Mayer during September and October, 1952,
when they travelled together up the Wahgi valley from Nondugl to Tomba. There
Mr. Loke compijed most of his field notes which he kindly sent to me ; and extracts
from most of them are included here. Two of his photographs are also published with
his kind permission.
I wish to acknowledge the generous co-operation of the authorities in the museums
at Berlin, Harvard, Leiden, New York and Pittsburgh. They have very kindly lent
me specimens for examination or made critical notes on specimens sent to them for
comparison. I would also like to acknowledge the help afforded to me by Mr. R. P.
Derek Goodwin. He made a preliminary examination of the Shaw-Mayer Collection
and I found his notes most useful. Finally I wish to thank Mr. J. D. Macdonald for
reading the manuscript and for his helpful criticism.
2. Methods
The report of the American Museum of Natural History Expeditions (Mayr and
Gilliard, 1954) contained descriptions of the topography and ecology of the area, and
lists of the birds typical of the various habitats and their zoogeographical affinities.
These subjects, therefore, receive no further treatment in this report except where
new information is presented.
The sequence of the families, genera and species and the nomenclature are that of
Mayr (19416). In that work a synonymy of each name is given and consequently
only the original reference to each name is normally given here.
The specimens are listed under their British Museum (Natural History) register
numbers together with information provided by the collector. The dimensions are
given in millimetres ; the wings were flattened when they were measured so the
measurement given is the arc ; the bill measurement is that of the exposed part of
the culmen.
BIRDS COLLECTED BY MR
SHAW-MAYER
39i
3. Summary of New Information
[a) Additions to the list of the birds of the Wahgi region (Gyldenstolpe, 1955).
Casaurius bennetti shawmayeri.
Dupetor flavicollis gouldi.
Falco peregrinus ernsti.
Porzana pusilla palustris.
Alcyone azurea ochrogaster.
Cisticola exilis diminuta.
Petroica bivittata bivittata.
Aplonis metallica metallica.
Neositta papuensis alba.
(b) Zoogeographical affinities of Wahgi region birds not listed by Mayr and Gilliard,
1954.
Southern affinities (Australia) : South-eastern affinities :
Porzana pusilla palustris Casuarius bennetti shawmayeri (Kratke
Mountains)
Cisticola exilis diminuta Circus spilonotus spilothorax
Petroica bivittata bivittata
North-eastern affinities (Huon Peninsula) :
Turnix maculosa giluwensis
Northern affinities (Sepik Region)
Alcyone azurea ochrogaster
Astrapia stephaniae femina
Western affinities (Mountains of Dutch New
Guinea) :
Neositta papuensis alba
(c) Habitats of Wahgi region birds not listed by Mayr and Gilliard, 1954.
Grasslands — 4,000 ft. to 7,500 ft. Mid-mountain forest — 7,500 ft. to
9,000 ft.
Dupetor flavicollis gouldi Casuarius bennetti shawmayeri
Anas super ciliosa rogersi Scolopax saturata rosenbergii
Circus spilonotus spilothorax Aplonis metallica metallica
Turnix maculosa giluwensis Astrapia stephaniae femina
A ley one azurea ochrogaster Neositta papuensis alba
Cisticola exilis diminuta
Timber-line, edge of the alpine grasslands — 11,000 ft.
Petroica bivittata bivittata
COLLECTING LOCALITIES
October, 1946 : Menebe, Mount Hagen, 5,000 ft.
October, 1950 : Headwaters of the Minj River, Kubor Range, 8,000 to 9,000 ft.
November, 1950 ; Minj, Wahgi River, 5000 ft. ; and the forested slopes of the
Kubor Range.
December, 1950 and January, 195 1 ; Forested ridges on the west slopes of the
Hagen Range near Welya, 7,500 ft. to 10,000 ft.
1 Racial identity problematical, Gyldenstolpe (1955).
2 Possibly includes Neositta papuensis wahgiensis Gyldenstolpe, 1955.
392
BIRDS COLLECTED BY MR. F. SHAW-MAYER
BIRDS COLLECTED BY MR. F. SHAW-MAYER 393
February and March, 1951 : South slopes of the Hagen Range near Tomba, 8,000
to 10,000 ft.
March to July, 1951 : Plateau grasslands north of Mount Giluwe, 7,300 ft. Lamende
Range north-west of Mount Giluwe, 8,500 ft. (23rd March). Moss Forest,
10,000 ft.; and the alpine grasslands, 11,000 ft.; Mount Giluwe (24~28th
April).
Moss Forest, 10,000 ft.; and the alpine grasslands, 11,000 feet, Lamende
Range (nth to 14th May).
Forested north slopes, 7,500 to 8,500 ft.; and the mid-mountain grasslands,
9,000 ft., Mount Giluwe (20th to 30th May).
Mount Giluwe rises to 13,414 ft. above sea-level from a grassland plateau of 7,300 ft..
The lower slopes of the mountain are forested up to about 9,000 ft. where the mid-
mountain grasslands commence. These are separated from the alpine grasslands of
the summit by the moss forest that extends approximately from 10,000 ft. to 11,000
ft.. The distribution of the forests and grasslands appears to be similar on the Lamende
Range and characteristic of the mountains of this area ; fuller details, particularly
of the Bismarck, Hagen and Kubor Ranges, were given by Mayr and Gilliard (1954 :
323).
ANNOTATED LIST OF BIRDS COLLECTED IN THE WAHGI REGION
BY MR. F. SHAW-MAYER
Casaurius bennetti shawmayeri Rothschild
Casuarius papuanus shawmayeri Rothschild, 1937, Bull. Brit. Orn. Club, 57, p. 120. Arau
district, Kratke Mountains, New Guinea.
1954.17.266 ; i $ ; Menebe, north of Mount Hagen, 5,000 ft.; October, 1946.
Measurements, i ? — height of back from ground, 860 ; total length, 1,170.
Colours : bill and casque, shiny black ; legs, horn ; iris, brown. Stomach Contents :
remains of hard fruits.
The specimen and a drawing of the head and neck at death agree with the type. The
record extends the known range of shawmayeri about 150 miles north-west from the
Kratke Mountains to the Hagen Range. It is probable that this race is found through-
out the Wahgi Valley area, but it remains to be seen how far it extends into the
mountains on each side. Mr. Gilliard saw a cassowary on Mount O'-Mar in the Kubor
range which he believed was this race (Mayr and Gilliard, 1954 : 331), and the present
specimen taken near Mount Hagen supports his provisional identification.
Dupetor flavicollis gouldi (Bonaparte)
Ardetta gouldi Bonaparte, 1855, Consp. Gen. Av. 2, p. 132. New South Wales, Australia.
1953. 17. i ; 1 (J ; Mount Giluwe, 7,300 ft.; June, 1951.
Measurements : 1 <$ — wing, 209 ; tail, 76 ; bill, 69. Colours ; bill, upper
mandible dark horn and lower mandible light horn ; legs, brown-grey ; iris, yellow.
Stomach Contents : water insects,
394 BIRDS COLLECTED BY MR. F. SHAW-MAYER
Anas superciliosa rogersi Mathews
Anas superciliosa rogersi Mathews, 1912, Austr. Av. Rec. 1, p. 33. Augusta, south-western
Australia.
1953. 17.2-4 ; i (J, 2 juv. cJ ; Mount Giluwe, 7,300 ft.; June, July, 1951.
Measurements : 2 juv. (J — wing, 229, 239 ; tail, 68, 71 ; bill 46, 48. 1 <J — wing
250 ; tail 96 ; bill 47. Colours : bill, slate (ad. <J black at base of lower mandible) ;
legs, pale brown flesh (ad. £ greenish pale brown, darker on webs of feet) ; iris,
brown. Stomach Contents : insect larvae, grass seeds and river gravel.
These birds were taken on the plateau north of Mount Giluwe where the species
was plentiful. The June birds appear to be juveniles because the feathers of the
undersurface have a dark brown streak unlike the adult, where they are dark brown
and fringed with cinnamon. The primaries are fully grown and show no signs of wear,
yet the wing is shorter than the adult's. In this respect the young birds approach the
smaller size of the other New Guinea race, pelewensis, which differs from rogersi only
in the shorter wing — -pelewensis, 230 ; rogersi ,250 — Amadon (1945 : 3) ; nevertheless,
this pair appear to be young birds of the larger form, rogersi. I have not seen
any material of comparable age and plumage, but it is likely that the primaries of
the juvenile plumage which persist throughout the first adult plumage are shorter
than the primaries of the full adult plumage. The young birds have started to moult
and a few adult patterned feathers have come in on the undersurface ; the wing does
not appear to be involved in the moult. The July bird, taken nearly six weeks later,
is completing moult. The feathers of the undersurface of all three specimens are
variably stained a rust colour, probably with iron oxide.
Salvadorina waigiuensis Rothschild and Hartert
Salvadorina waigiuensis Rothschild and Hartet, 1894, Novit. Zool. 1, p. 683. ? Waigeu, possibly
Vogelkop, north western New Guinea.
1953. 17.5-6 ; 2 ? ; Mount Giluwe, 7,300-7,200 ft.; April, July, 1951.
Measurements : 2 °- — wing, 180-185 ; tail, 95 ; bill, 34-35. Colours : bill, dull
yellow with black along the centre of the upper mandible ; legs, yellowish horn (webs
of feet almost black) ; iris, dark brown. Stomach Contents : river gravel, water
beetles and other small aquatic insects.
Accipiter melanochlamys schistacinus (Rothschild and Hartert)
Astur melanochlamys schistacinus Rothschild and Hartert, 191 3, Novit. Zool. 20, p. 482. Mt.
Goliath, Oranje range, central New Guinea.
1953. 17.7-8 ; i (J, i $ ; Mount Giluwe ; 8,000 ft.; May, July, 1951.
Measurements : 1 <J — wing, 216 ; tail, 160 ; bill, 15-5. 1 $ — wing, 251 ; tail,
184 ; bill, 18. Colours : bill, black (cere — $ yellow and $ orange-yellow) ; legs,
BIRDS COLLECTED BY MR. F. SHAW-MAYER 395
J bright yellow and ? orange ; claws, black ; iris, <J bright yellow and $ bright orange.
Stomach contents : £, remains of small bird ; ?, remains of frog.
The female (July) is in moult and the new black feathers of the back have a
greenish metallic lustre in contrast to the worn feathers which have faded to a rusty
brown-black. The male (May) is a young bird in fresh immature plumage. The
exposed parts of the feathers of the back are chestnut in colour edged with rufous brown,
the unexposed parts being white. The feathers of the shoulders and most of the head
are white tipped with black, while the crown is chestnut in colour. The primaries and
the retrices are dark brown with black-brown barring above ; they are silvery below
with irregular cinnamon coloured areas, particularly on the inner webs. The feathers
of the under-surface are cream in colour with a median chestnut stripe while those of
the legs are more buffy with a narrow rufous stripe.
Circus spilonotus spilothorax Salvadori and D'Albertis
Circus spilothorax Salvadori and D'Albertis, 1875, Ann. Mus. Civ. Genova, 7, p. 807. Yule
Island, south-eastern New Guinea.
I 953-i7-9 ; 1 c? ; Mount Giluwe, 7,300 ft.; July, 1954.
Measurements : 1 $ — wing, 375 ; tail, 220 ; bill, 21-5. Colours : bill, black
(cere, bluish horn ; dorsal surface, greenish yellow.) ; legs, dull yellow ; claws,
black ; iris, bright yellow ; inside of mouth, blue-grey.
" I often observed this fine bird soaring over the grasslands of the plateau, some-
times high in the air, at other times a few feet above the grass. Several times it came
to rest on the ground, but it was not observed to be eating anything. At the approach
of the hunter it was off, sailing with speed in ever widening circles. Once noticed at
dusk, but more often during the early morning. It appeared to be alone." (F. S.-M.).
In comparison with other birds of this race this specimen is considerably darker in
colour, both above and below. The upper-parts are a rusty black and lack the typical
silvery white wash which is present only over the upperside of all except the inner
and outer pairs of rectrices. The inner rectrices are almost black above and the outer
a faded chestnut colour, none being spotted like those of other specimens. This bird
exhibits a type of melanism not uncommon among birds of prey, but it appears to be
darker than any reported by Rand (1941a : 1).
Falco peregrinus ernesti Sharpe
Falco ernesti Sharpe, 1894, Ibis, Ser. 6, 4 : 545. Mt. Dulit, Sarawak.
^S-^-io-; icJ; Mount Giluwe, 7,3000 ft.; July, 195 1.
Measurements : 1 <$ — wing, 292 ; tail, 144 ; bill, 19. Colours : bill, dark horn
around cere and gape, base of mandible yellow ; legs, yellow (claws, black) ; iris,
black (skin around eye, yellow).
396
BIRDS COLLECTED BY MR. F. SHAW-MAYER
Aepypodius arfakianus (Salvadori)
Talegallus arfakianus Salvadori, 1877, Ann. Mus. Civ. Genova, 9 : 333. Arfak Mountains, north-
western New Guinea.
1953.17.11-12 ; i (J, i ? ; Mount Giluwe, 9,000 ft.; April, July, 1951.
Measurements : 1 <J — wing, 278 ; tail, 132 ; bill, 31. 1 ? — wing, 272 ; tail, 145 ;
bill, 30. Colours : bill, yellowish-green-horn ; legs, greenish-yellow ($, frontal
scutes, dark horn) ; iris, greenish yellow ; head, pale bluish white, reddish colouring
along top of gape from back of beak to below eye (<?, purple comb and wattles,
bluish area on crown to hinder end of the base of the comb ; $, two small pinkish
wattles — no comb — bluish area above wattles). Stomach contents : pieces of hard
fruits.
The birds were taken in the forest on the north slopes of Mount Giluwe. An egg
was found in the oviduct of the female (July).
Synoicus ypsilophorus lamonti Mayr and Gilliard
Synoicus ypsilophorus lamonti Mayr and Gilliard, 195 1, Amer. Mus. Novit. No. 1524, p. 1.
Central Highlands, New Guinea.
Tomba
1953 . 17 . 13-24 ; 7 c?, 5 ? ; Mount Giluwe, 7,500-7,200 ft.; April, May, 1951.
Measurements : 7 $ : wing, 90-100 ; bill, 12-13-5. 5 ? — wing, 93-98 ; bill, 12-13.
Colours ; bill, black, bluish horn at base ; legs, greenish yellow ; iris, yellow-orange.
Stomach contents : chiefly grass seeds, some insects and gravel.
Most of the specimens of this fine series match closely 2 (J and 2 $ of the race
lamonti taken by Mr. Gilliard at Tomba, which is only a few miles north of Mount
Giluwe. The exception is the male bird (No. 1953. 17. 19) taken on 27th May. In
general colour it is a much paler brown, more heavily barred above and below and the
white shaft streaks of the feathers of the back are wider and more pronounced. It
was sent to the American Museum of Natural History where, in Mr. Gilliard's absence,
Dr. Amadon kindly compared it with other material of 5. ypsilophorus. He found
that it was unlike any other bird of this species and that the differences between this
specimen and other examples of lamonti are greater than those between lamonti and
neighbouring races. As these races appear to be fairly stable in their characteristics
we have concluded that this specimen is probably aberrant.
" This quail was plentiful on the grasslands at 7,000 feet, and on the grasslands on
the north slopes of Mount Giluwe up to 8,000 feet. Sometimes a single bird would flush,
at other times three or more. Much of the country is peaty, almost swampy, and these
conditions seem to suit it best " (F. S.-M.).
Field Notes. Six of these quail were started up in pairs from the long grass near
Nondugl. A roughly made nest of grass was found on the ground among the weeds
near some land under cultivation on 15th September. The nest contained four
buff-coloured eggs with thickly distributed minute brown spots.
BIRDS COLLECTED BY MR. F. SHAW-MAYER 397
Excalfactoria chinensis subspecies ?
Tetrao chinensis Linnaeus, 1766, Syst. Nat. 12th ed., 1, p. 277. China.
1953.17.25-33 ; 3 cj, 6 ? ; Mount Giluwe, 7,500-7,200 ft.; April, May, 1951.
Measurements : 3 <J — wing, 65-68 ; bill, 8-9. 6 $■ — wing, 65-70 ; bill, 9-10.
Colours : bill, black ; legs, orange, orange-yellow (2 $ with greenish tinge) ; iris,
3 c? — brownish-red ; 6 ? — dark brown. Stomach Contents : grass seeds and gravel.
The series was collected on the grasslands north of Mount Giluwe. The birds are in
fresh plumage with the exception of the female taken on 7th April which is in moult
although it had an egg in the oviduct. Two races of E. chinensis have been described
from New Guinea, papuensis Mayr and Rand from Mafulu in the south-east and
novaeguineae Rand from the Balim River in the Snow Mountains. Mount Giluwe is
approximately midway between the type localities of these races and, as might be
expected, the birds from there exhibit characteristics of both. The series was com-
pared with specimens of both races collected by Dr. Rand. The adult males differ
but little from a male taken at Lake Daviumbo on the Fly River and compared with
typical papuensis the bluish colour of the head is slightly less pronounced and the
small feathers of the head have dark grey-brown tips ; in this respect they approach
the characteristics of novaeguineae. The adult female from Mount Giluwe differ
from papuensis only in the colour of the under-parts, where there are gradations from
the richer colour of papuensis to the paler colour of novaeguineae.
The series is almost intermediate between papuensis and novaeguineae. I am
not, therefore, naming them racially. Furthermore, Dr. Rand's birds are in old
plumage, whereas in most of these the plumage is fresh.
Turnix maculosa gilutvensis Sims
Turnix maculosa giluwensis Sims, 1954, Bull. Brit. Orn. Club, 74, p. 37. Mount Giluwe, Central
Highlands, New Guinea (8,000 ft.).
I 953-i7-34-4i 1 ; i(J,i juv. $; 3 <j>, 3 juv. $ ; Mount Giluwe, 8,000-7,200 ft.;
May, June, 1951.
Measurements: 2 $ — wing, 65; tarsus, 18. 5-19; bill, 10-n. 6 ? — wing,
70-75 ; tarsus, 19-20 ; bill, 11-5-14. Colours : bill, yellowish, ridge and tip dark
horn; feet, yellow with greenish tinge; iris, cream. Stomach contents: grass
seeds and small insects.
This fine series of Button-quail was taken on the open and lightly wooded grass-
lands around Mount Giluwe. It appears that this is the first time that such a large
series of T. maculosa has been taken in one locality in New Guinea ; the males,
particularly, are rare to collections. On examination these birds were found to be
distinct enough to recognize as representatives of a new race of T. maculosa in
1 Type 1953.17.40
398 BIRDS COLLECTED BY MR. F. SHAW-MAYER
comparison with the other two New Guinea races, furva Parkes from Finschhafen
on the east coast of the Huon Peninsula and horsbrughi from Yule Island off the
south-east coast. They differ from these races by the underparts being considerably
paler, but resemble furva by being very dark above. In size giluwensis is closer to
horsbrughi than to furva, which is slightly larger. The one June bird is in fresh
plumage the others are involved in moult.
Rallus pectoralis captus Mayr and Gilliard
Rallus pectoralis captus Mayr and Gilliard, 1951, Amer. Mus. Novit. No. 1524, p. 2. Tomba,
Central Highlands, New Guinea.
1953.17.42 ; i(J; Hagen Range, 8,000 ft.; February, 1951.
1953.17.43-45 ; 1 (J, I juv. (?, 1 $ ; Mount Giluwe, 7,500-7,300 ft.; April, June,
Measurements : 3 <J — wing, 102-104 ; tail, 37-40 ; bill, 33. 1 $ — wing, 102 ;
tail, 40 ; bill, 34. Colours : bill, dull red, dark horn near tip ; legs, brownish grey ;
iris, yellow to brown (juv. <J, grey brown). Stomach contents : slugs and water
insects.
The adult male and female taken in February and April respectively are in worn
plumage ; the barring of the abdomen is faded and the flanks are stained a pinkish
hue. The adult male taken in June is in fresh plumage ; but the young male, also
taken in June, is in full moult. In this bird the new feathers of the under-parts, unlike
the worn ones, are strongly barred like those of the adults. The adult male from the
Hagen Range is topotypical and both it and the other adult male show the character-
istics of captus given by Mayr and Gilliard, but the wings average a little longer
(wing, <$ 95-102), like those of specimens taken at Nondugl by Gyldenstolpe (1955 :
32). The adult female, in comparison with the adult male, has the back paler in
colour by being a clearer olive-brown, less reddish. In the juvenile male the crown
and the neck lack the maroon-chestnut of the adult and there is a reduction in the
white barring of the back. In this respect the young bird agrees with a series of
adults of R. p. pectoralis (Temminck) from eastern Australia.
Rallus philippensis wahgiensis Mayr and Gilliard.
Rallus philippensis wahgiensis, Mayr and Gilliard, 1951, Amer. Mus. Novit. No. 1524, p. 3.
Nondugl, Central Highlands, New Guinea.
1953.17.46 ; i <$ ; Mount Giluwe ; 7,500 ft.; July, 195 1.
Measurements : 1 $ — wing, 153 ; tail, 76 ; bill, 31 ; tarsus, 41. Colours : bill,
dark horn, base of lower mandible reddish ; legs, light brown ; iris, reddish brown.
Stomach contents : insects.
This specimen differs only in size from a topotypical male collected by Mr. Gilliard.
The wing and the tail are longer and the tarsus shorter, the measurements of
BIRDS COLLECTED BY MR. F. SHAW-MAYER 399
wahgiensis given by Mayr and Gilliard are : wing, 137, 140 ; tail, 62.5 ; tarsus, 44,
45. In this respect it matches specimens taken at Nondugl by Gyldenstolpe (1955 : 32) .
Porzana pusilla palustris Gould
Porzana palustris Gould, 1843, Proc. Zool. Soc. Lond. 1842, p. 139. Tasmania.
Porzana pusilla mayri Junge, 1953, Zool. Meded., Leiden, 31, p. 247. Paniai, western New
Guinea.
1953.17.47-49 ; 2 ?, i juv. $ ; Mount Giluwe, 8,000-7,500 ft.; May, June, 1951.
Measurements : 3 $ — -wing, 79-83 ; tail, 35-40 ; bill, 14-3-15. Colours : bill,
olive-green, tip horn colour ; legs, olive-green ; iris, orange-red. Stomach contents :
small insects.
The Mount Giluwe birds may be distinguished from a series of palustris from
eastern Australia on very slight colour differences. The upper-parts of the New
Guinea birds are somewhat warmer in tone and the under-parts a slightly clearer
grey, but these differences may be attributable to the freshness of the skins. Junge
(1952 : 247) was the first to report the presence of this species in New Guinea. He
distinguished a new race, mayri, on a series from Paniai separating it from the Austra-
lian birds on a greater wing length. The measurements of the material before me,
however, do not support this separation for there is some variation.
Australian birds, in B.M. (N.H.)
„ (Junge, 1952 : 247)
New Guinea ,, in B.M. (N.H.)
„ „ (Junge, 1952 : 247)
Wing, 75-81 1
„ 80-85/ 75 ° 5
" T 83 >76-8 3
„ 76-79/ / J
Porzana tabuensis edwardi Gyldenstolpe
Porzana tabuensis edwardi Gyldenstolpe, 1955, Ark. f. Zool., Ser. 2, 8, p. 3. Nondugl, Central
Highlands, New Guinea.
1953.17.50-56; 2 (J, i juv. (J ; 2 ?, 2 juv. ?; Mount Giluwe, 8,000-7,5000 ft.;
March, April, June, 195 1.
Measurements : 3 <$ — wing, 78-85 ; tail, 41-50 ; bill, 18-19. 4 ¥ — wing,
78-80 ; tail, 41-46 ; bill, 16-18. Colours : bill, black ; legs, coral-red ; iris, orange-
red. Stomach contents : insects and grass seeds.
Gyldenstolpe (1955 : 34) separated the birds of the Wahgi Valley on their darker
colour and slightly longer bill in comparison with a series of tabuensis from the Tuamoto
Archipelago and Rapa Island. These characters are present in the Mount Giluwe
birds and distinguish them from a series from various Pacific islands. The adult male
taken in April is in worn plumage, while that taken in June is in fresh. The fresh
plumage is more vinous above and more blue than slate-grey below. The young male,
31st March, differs from the adult in having the chin and throat white and the
4 oo BIRDS COLLECTED BY MR. F. SHAW-MAYER
abdomen and under- wing coverts a pale whitish-grey. This bird is in moult. The adult
females taken in April are starting moult with a few new feathers coming through on
the back. Of the two young females one, 5th April, has nearly completed moult ;
while the other, 7th April, has only a few new feathers coming through on the back.
Rallicula forbesi subspecies?
Rallicula forbesi Sharpe, 1887, in Gould's "Birds of New Guinea", pt. 23, pi. (70). Owen Stanley
Range, south-western New Guinea.
1953.17.57 ; i(J; Hagen Range ; 8,000 ft.; February, 1951.
1953 . 17 . 58-62 ; 1 (J, 4 $ ; Mount Giluwe ; 7,400-7,300 ft.; May, June, July, 1951.
Measurements : 2 $ — wing, 111-112 ; tail, 77-79 ; bill, 24-25. 4 $ — wing,
108-112 ; tail, 63-68 ; bill, 24-26. Colours : bill, brown-black ; legs, brown-black
to black ; iris, pale to dark brown.
The two males differ only very slightly from the type and a series of forbesi from
south-eastern New Guinea in that the plumage is somewhat richer in tone. The four
females, on the other hand, exhibit an interesting variation. Although they appear to
be inseparable from the allotype and a series of forbesi in most respects there are
differences in the speckling and the occurrence of a red-chestnut fringe on the feathers
of the back. The July bird agrees with the allotype in colour and pattern in that over
the back the speckling is the typical buff colour and many of the feathers are fringed
imperfectly with a red-chestnut colour. Gradations are found in the other females to
the colour and pattern of a May bird (1953. 17.62) in which the spotting is no longer
buff but white, and replaced over the scapulars by white streaks ; also none of the
feathers of the back have a red-chestnut fringe. The markings of this specimen so
approached the description of the race steini which Rothschild described on one female
from the Weyland Mountains that it was sent to the American Museum to be compared
with the unique type. There Dr. Amadon, in Mr. Gilliard's absence, kindly examined
the specimens and found that the Mount Giluwe bird matched the type. Mayr and
Gilliard (1954 : 335) described a similar variation in a Wahgi region series which they
named steini mainly on the length of the tail. The present series, however, cannot be
separated from the nominate form on measurement, while on plumage most of the
birds approach forbesi. It seems that if the present series were combined with the
birds reported by Mayr and Gilliard then phenotypical forbesi and steini would be
present in approximately equal numbers in addition to the specimens grading
between the two. In view of this I do not venture to name the birds of the present
series at racial level.
One explanation of the occurrence of this variation in these series from the Wahgi
region could be that the two races have come into breeding contact following a period
of reproductive isolation. It seems that such an interpretation, based on the material
available, would not be entirely incorrect because a population showing such a high
incidence of heterogeneity would be unlikely to occur in a cline. It seems equally
BIRDS COLLECTED BY MR. F. SHAW-MAYER 401
unlikely that the type of steini and other specimens that resemble it are variants for
no similarly patterned birds have been reported from southeast of the Wahgi region.
Pluvialis dominca fulva (Gmelin)
Charadrius fulvus Gmelin, 1789, Syst. Nat. 1, pt. 2, p. 687. Tahiti.
Field Note. Six of these plovers were seen together in a " paddock " at Nondugl
on 15th September. A few of the black feathers of the breeding plumage could be
seen on the under-surface of some of the birds.
Scolopax saturata rosenbergii Schlegel
Scolopax rosenbergii Schlegel, 1871, Ned. Tijdschr. Dierk. 4, p. 54. Arfak Mts., north-western
New Guinea.
1953.17.63-65; 2 <J, 1 ? ; Mount Giluwe, 9,500-9,000 ft.; May, June, 1951.
1953.17.66-67 ; 1 (J, 1 ? ; Lamende Range, 9,500-9,000 ft.; June, 1951.
Measurements : 2 <J — wing, 145-146 ; tarsus, 33-34 ; bill, 82-83. 3 $ — wing,
148-155 ; tarsus, 32-35 ; bill, 83-87. Colours : bill, brown-black horn ; legs,
dark brown-grey ; iris, dark brown. Stomach contents : worms, grubs, small
beetles and other insects.
" The Woodcock is a bird of the high mountain-forest on Mount Giluwe and the
Lamende Range. It lives on the ground, where it searches for insects and digs for
worms and grubs. During the early morning it will often rise and fly high above the
trees, where it can be heard calling ; later it drops down again on to the forest floor ' '
(F. S.-M.).
The black barring of the feathers of the back is appreciably heavier than in topo-
typical specimens of rosenbergii. In this respect the present series is similar to the
specimens that I have seen from the Oranje Mountains taken by Dr. Rand who
considered that this difference was of no racial significance. (19426 : 439). The wing
measurements are less that those of Dr. Rand's specimens but Shaw-Mayer's birds are
in moult and it seems that the primaries are not fully grown. In these moulting birds
the brown of the new feathers is noticeably redder and darker than that of the old.
Gallicolumba beccarii beccarii (Salvadori).
Chalcophaps beccarii Salvadori, 1875, Ann. Mus. Civ. Genova, 7, p. 974. Hatam, Arfak Moun-
tains, north-western New Guinea.
1953.17.68 ; i (J ; Hagen Range, 8,000 ft.; March, 1951.
1953.17.69 ; 1 c? ; Mount Giluwe, 7,500 ft.; May, 1951.
Measurements : 2 ad. <J — wing, 107,109 ; tail, 60,70 ; bill, 14. Colours : bill,
black ; legs, dark purple ; iris, brown-black. Stomach contents : hard seeds.
4 02 BIRDS COLLECTED BY MR. F. SHAW-MAYER
Charmosyna papou goliathina Rothschild and Hartert
Chavmosyna stellae goliathina Rothschild and Harter, 191 1, Novit. Zool. 18 ,p. 160. Mt. Goliath,
Oranje Range, central New Guinea.
1953 . 17 . 70-73 ; 2 (J, 2 $ ; Mount Giluwe, 9,000-8,500 ft.; April, June, July, 1951.
Measurements : 2 (J— wing, 151, 155 ; tail, 297, 324 ; bill, 18. 2 ? — wing, 150,
152 ; tail, 256, 280 ; bill, 17, 19. Colours : bill, orange to dark red ; legs, bright
orange ; iris, orange-red. Stomach contents : flowers and fruit pulp.
The two males are in the dark phase plumage. The June and July birds of both
sexes are completing moult.
Field note. Native name, " Kai-niginch ".
Oreopsittacus arfaki grandis Ogilvie-Grant
Oreopsittacus grandis Ogilvie-Grant, 1895, Bull. Brit. Orn. Club, 5, p. 15. Owen Stanley
Mountains, south-eastern New Guinea.
1953.17.74-76 ; i <J, 2 ? ; Hagen Range, 9,500-9,000 ft.; December, 1950.
Measurements : 1 <J — wing, 81 ; tail, 84 ; bill, 8. 2 $ — wing, 82, 86 ; tail, 85 ;
bill, 8, 9. Colours : bill, black ; legs, green-grey ; iris, black-brown.
These birds match grandis except in wing length. In this respect the females are
larger than both grandis and major and the males intermediate, — grandis : $, 73-76 ;
9, 73 — major : (J, 85-87 ; $, 80 — (Ogilvie-Grant, 1915 : 169).
Neopsittacus musschenbroekii major Neumann
Neopsittacus musschenbroekii major Neumann, 1924, Ornith. Monatsber. 32, p. 38. Schrader-
berg, Sepik Mountains, northern New Guinea.
I953-I7-77 ; 1 (J ; Mount Giluwe, 7,500 ft.; June, 1951.
Measurements : 1 <$ — wing, 114 ; tail, 101 ; bill, 16. Colours : bill, yellow ;
legs, brown-grey ; iris, orange-red. Stomach contents : fruits and berries.
Other birds taken in the same area were identified as major by Mayr and Gilliard
(1954 : 339) but the present example differs from other material of the species in
being much darker. Compared with major, the crown, collar, nape and sides of the
throat are browner and washed with crimson. The chin and throat are dark brown
with a greenish wash. The back appears to be very dark due to the feathers being
fringed with dark brown. Some of the primary wing coverts, the outer webs of the
primaries and the upper surfaces of the two central rectrices are almost entirely dark
brown with a slight greenish wash. The pattern of the underparts is similar to major,
but the green is replaced by olive and the crimson by a yellowish-brown. This
specimen may be an abnormally pigmented bird of the race major, although the
possibility that it represents a distinct form should not be overlooked.
BIRDS COLLECTED BY MR. F. SHAW-MAYER 403
Psittacella brehmii pallida A. B. Meyer
Psittacella pallida A. B. Meyer, 1886, Zeitschr. ges. Ornith. 3, p. 3. Astrolabe Mountains,
south-eastern New Guinea.
Psittacella biirgersi Reichenow, 1918, Journ. f. Ornith. 66, p. 244. Schraderberg, Sepik region,
northern New Guinea.
Psittacella brehmi ornata Mayr, 193 1, Mitt. Zool. Mus. Berlin, 17, p. 720. Dawong, Herzog
Mountains, south-eastern New Guinea.
1953.17.78-79 ; i <J, i $ ; Mount Giluwe, 8,500 ft.; June, 1951.
1953. 17.80-81 ; 1 (J, 1 juv. $ ; Hagen Range, 9,000 ft.; February, 1951.
Measurements : 2 <$ — wing, 122, 125 ; tail, 86 ; bill, 18, 20. 1 ? — wing, 127 ;
tail, 87 ; bill, 18. 1 juv.? — wing, 122 ; tail, 86 ; bill, 17. Colours : bill, metallic
blue-grey, becoming white towards tip in juvenile ; legs, dark grey, green-grey in the
ad. (J ; iris, orange, grey in juvenile. Stomach contents : hard berries and seeds.
Gyldenstolpe (1955 : 57) described much of the individual variation in both color-
ation and size in the birds of this species from central and south-eastern New Guinea.
He showed that biirgersi should be regarded as a synonym of pallida and agreed with
Mayr (1941 : 69) who had previously united ornata with pallida. Of the present
specimens the adult female and the juvenile were sent to the Berlin Museum, where
Professor Stresemann, who kindly made the comparison, found that they closely
matched the type of P. biirgersi Reichenow. In view of the evidence presented by
Gyldenstolpe I am naming the birds of this present series pallida. Compared with a
small series of intermedia from southern Dutch New Guinea the Wahgi region birds
are more bluish-green above with a greenish wash over the crown ; and the sides of
the head are more greyish in colour.
Psittacella picta excelsa Mayr and Gilliard
Psittacella picta excelsa Mayr and Gilliard, 1951, Amer. Mus. Novit. No. 1524, p. 6. Mount
Orata, Kubor Mountains. Central Highlands, New Guinea.
1953.17.82 ; I (J ; Hagen Range, 9,000 ft.; February, 1951.
1953.17.83 ; 1 juv. $ ; Kubor Range, 8,000 ft.; October 1950.
1953.17.84 ; 1 (J ; Mount Giluwe, 11,000 ft.; April, 1951.
Measurements : 2 <J — wing, 114, 116 ; tail, 73 ; bill, 14, 15. 1 juv. $ — wing,
103 ; tail, 68 ; bill, 12-5. Colours : bill, pale blue, yellowish (juv. (J) ; legs, grey ;
iris, orange-yellow, grey-yellow (yg. cJ). Stomach contents : seeds and small hard
berries.
The adult from the Hagen range differs from the Mount Giluwe bird and a topo-
typical specimen taken by Mr. Gilliard in that the golden-yellow patches on each
side of the neck are joined over the neck by a narrow band of the same colour. The
young male resembles a topotypical female also collected by Mr. Gilliard.
ZOOL. 3, IO. 26
4°4
BIRDS COLLECTED BY MR. F. SHAW-MAYER
Psittacella modest a hallstromi Mayr and Gilliard
Psittacella modesta hallstromi Mayr and Gilliard, 1951, Amer. Mus. Novit. No.
Yandarra, north slope of Mount Wilhelm, Bismarck Mountains, New Guinea.
1524,
P- 5
1953.17.85-87 ; 2 (J, 1 $ ; Mount Giluwe, 8,500-8,000 ft.; April, June, 1951.
1953.17.88 ; 1 juv. $ ; Rubor Range, 8,000 ft.; October, 1950.
1953.17.89 ; 1 <J ; Hagen Range, 9,500 ft.; February, 1951.
Measurements : 4 <J — wing, 98-101 ; tail, 59-62 ; bill, 14. 1 ? — wing, 102 ;
tail, 64 ; bill, damaged. Colours : bill, metallic blue-grey ; legs, dark blue-grey ;
iris, orange (2 ad. (J), yellow-brown (2 imm. (J), dark brown (1 ad. ?). Stomach
contents : seeds and pieces of hard berries.
The young male taken in October is starting to moult from juvenile plumage in
which the throat and breast are an olivaceous colour and lack the ashy-brown of the
adult male. The feathers of the breast are also faintly barred with yellow as in the
female where the orange and black barring of the breast merges into the plain green
of the abdomen and flanks. The head and the nape are the same colour as the back
although the feathers at the base of the bill are washed with blue and some on the
hind neck are heavily barred with orange and black. The February bird is in worn
plumage, while the April and June birds are in fresh plumage. The testes of the June
male are enlarged.
Caco mantis pyrrhophanus excitus Rothschild and Hartert
Cacomantis excitus Rothschild and Hartert, 1907, Novit. Zool. 14, p. 436. Angabunga River,
south-eastern New Guinea.
1953. 17.90-91 ; i (J, i juv. (J ; Mount Giluwe, 7,500 ft.; April, May, 1951.
Measurements : 1 $ — wing, 145 ; tail, 140 ; bill, 17. 1 juv. <J — wing, 122 ; tail,
100 ; bill, 14. Colours : bill, black ; legs, yellow ; iris, dark brown. Stomach
contents : caterpillars and flying insects.
These birds were taken in the wooded country on the north slopes of Mount Giluwe.
The May bird is similar to a male from the Utakwa River in Dutch New Guinea
except that the grey of the chin extends to the throat. The young male which is in
fresh plumage (April) is plain chestnut above and rufous-brown irregularly barred
with black below.
Chalcites meyerii (Salvadori)
Chrysococcyx meyerii Salvadori, 1874, Ann. Mus. Civ. Genova, 6, p. 82. New name for Chryso-
coccyx splendidus A. B. Meyer, 1874, Sitzungsber, Akad. Wiss. Wien, 69, p. 81. Hatam,
Arfak Mountains. (Not Gray, 1847).
1953.17.92 ; 1 c? ; Wahgi River ; 5,000 ft.; November, 1950.
Measurements : 1 ? — wing, 88 ; tail, 65 ; bill, 12-5. Colours
legs, blue-grey ; iris, dark brown.
bill, black
BIRDS COLLECTED BY MR. F. SHAW-MAYER 405
Tyto tenebricosa arfaki (Schlegel)
Strix tenebricosa arfaki Schlegel, 1879, Notes Leyden Mus. 1, p. 101. Arfak Mountains, north-
western New Guinea.
1953.17.95 ; i (J ; Hagen Range, 8,500 ft.; March, 1951.
1953.17.96 ; 1 ? ; Lamende Range, 8,500 ft.; March, 1951.
Measurements : 1 $ — wing, 275 ; tail, 119 ; tarsus, 62. 1 ? — wing, 292 ; tail,
130 ; tarsus, 73. Colours : bill, dark horn (ad. $, lower mandible pale horn) ; legs,
dark flesh ; iris, brown-black. Stomach contents : 1 (J, remains of a rat.
Tyto longimembris papuensis Hartert
Tyto longimembris papuensis Hartert, 1929, Novit. Zool. 35, p. 103. Owgarra, Angubunga
River, south-eastern New Guinea.
1953.17.93-94 ; 1 <J, 1 $ ; Mount Giluwe, 7,400 ft.; May, 1951.
Measurements : 1 ad. $ — wing, 343 ; tail, 114 ; tarsus, 84. 1 ad. ? — wing, 357 ;
tail, 123 ; tarsus, 97. Colours : not recorded. Stomach contents : nil.
" <5, taken from nest in tree-hollow brooding three well incubated eggs ; time,
10 a.m. $, taken near hollow containing nest ; time, 10 a.m. Usually one or two of
these Grass Owls could be seen flying low over the grasslands or sitting on an old tree
fern stump during the late afternoon or at dusk. The species was, however, uncommon
on the plateau, possibly the altitude was a little too great for it. It was plentiful on
the grasslands east of the Hagen Range at about 5,000 feet." (F. S.-M.).
It is interesting that the male was taken while brooding the eggs as it is generally
accepted that it is not usual for the males of this species to share in incubation. The
eggs were " well incubated " on the 26th May, so it appears that the breeding season
of the New Guinea bird is similar to that of walleri, from Australia, which breeds
during May and June. It is worth mentioning, perhaps, that in November, 1946, Mr.
Shaw-Mayer collected a pair of Grass Owls at Yanka and in comparison the birds
from Mount Giluwe are slightly more tawny above and none of the feathers of the
back have white shaft streaks. The male from Yanka resembles the females in that
the feathers of the sides of the face-ring and the chin are tipped with brown and the
face is washed with the same colour, whereas these parts are white in the male
from Mount Giluwe.
Ninox theomarcha theomarcha (Bonaparte)
Spiloglaux theomarcha Bonaparte, 1885, Compt. Rend. Acad. Sci. Paris, 41, p. 654. Triton
Bay, south-western New Guinea.
1953.17.97 ; 1 juv. $ ; Hagen Range, 8,200 ft.; February, 1951.
Measurements : 1 juv. $ — wing, 188 ; tail, 98 ; tarsus, 29. Colours : bill, dark
horn ; legs, pale green-flesh ; iris, bright yellow. Stomach contents : remains of
large insects.
4 o6 BIRDS COLLECTED BY MR. F. SHAW-MAYER
The bird has started post-juvenile moult with new feathers coming through on the
breast, but towards the vent the paler juvenile plumage persists.
Podargus papuensis Quoy and Gaimard
Podargus papuensis Quoy and Gaimard, 1830, Voy. Astrolabe, Zool. 1, p. 207 (pi. 13). Dorey,
Vogelkop, north-western New Guinea.
1953.17.98 ; 1 $ ; Mount Giluwe, 7,300 ft.; May, 1951.
Measurements : 1 ad. ? — wing, 285 ; tail, 271 ; tarsus, 20. Colours : bill,
horn ; legs, dark grey ; iris, dark orange-red. Stomach contents : beetles.
Taken in the woodlands at the north base of Mount Giluwe.
Aegotheles albertisi salvadori Hartert
Aegotheles salvadori Hartert, 1892, Cat. Bds. Brit. Mus. 16, p. 649. Astrolabe Mountains,
south-eastern New Guinea.
1953 . 17 . 99-103 ; 2 (J, i $, 2 juv. ? ; Hagen Range, 8,500 ft.; February, 1951.
1953 . 17 . 104-105 ; 1 juv. (J, 1 juv. $ ; Mount Giluwe, 7,500 ft.; April, May, 1951.
Measurements : 3 <$ — wing, 118-121 ; tail, 97-98 ; bill, 8-9. 4 $ — wing,
116-125 ; tail, 97-102 ; bill, 8-9. Colours : bill, upper mandible dark horn and
lower mandible pale horn ; legs, dark flesh ; iris, dark brown. Stomach contents :
insects.
Nearest to salvadori but like the specimens described by Mayr and Gilliard
(1954 : 342) they differ from the typical form by the underparts being somewhat more
coarsely marked. The younger birds are moulting, the older are mostly in worn
plumage.
Aegotheles insignis insignis Salvadori
Aegotheles insignis Salvadori, 1875, Ann. Mus. Civ. Genova, 7, p. 916. Hatam, Arfak Mountains,
north-western New Guinea.
1953. 17. 106 ; i c? ; Mount Wilhelm, Bismark Range ; 6,000 ft.; May, 1951.
1953.17.107-111 ; 2 <J, 1 juv. cJ, 3 $ ; Hagen Range, 8,500-8,000 ft.; February,
March, 195 1.
1953.17.112-114 ; 1 juv. (J, 2 ? ; Mount Giluwe, 8,000-7,500 ft.; March, June,
I95i.
Measurements : 4 <$ — wing, 157-170 ; tail, 132-142 ; tarsus, 21-22. 5 °. — wing,
158-177 ; tail, 135-155 ; tarsus, 21-23. Colours : bill, upper mandible dark
brown and lower mandible pale horn ; legs, flesh ; iris, dull yellow to brown.
Stomach contents : beetles and other insects.
BIRDS COLLECTED BY MR. F. SHAW-MAYER 407
This series exhibits the colour phases of this race reported by Hartert (1930 : 95)
and Rand (19426 : 456). There are three light phase and six dark phase birds, three
of the latter being considerably darker than any I have seen before.
Eurostopodus archboldi (Mayr and Rand)
Lycornis archboldi Mayr and Rand, 1935, Amer. Mus. Novit. No. 814. p. 4. Mount Tafa,
Wharton Range, south-eastern New Guinea.
1953. 17. 119 ; i ^ ; Tomba, Hagen Range, 8,000 ft.; February 1951.
1953.17.120-121 ; 2 $ ; Mount Giluwe, 7,500-7,300 ft.; May, June, 1951.
Measurements : 1 $ — wing, 120 ; tail, 156 ; bill, 9-5. 2 $ — wing, 123, 125 ;
tail, 145, 149 ; bill, 10. Colours : bill, black ; legs, dark grey ; iris, brown-black.
Stomach contents : insects.
The male is in worn plumage and there is little spotting on the breast because many
of the feathers have lost their rufous tips. The May female is in fresh plumage and
shows numerous breast spots. A further difference between these specimens seems
to be due to fading for the colour of these spots is paler in the male than in the female.
These differences are interesting because they are qualitatively the same but quanti-
tatively greater than those between two specimens collected by Dr. Rand ; namely,
a male taken at Lake Habbema in Dutch New Guinea and a topotypical female
taken in September. These match the male and the May female respectively of the
present series, even in the plumage of the male being more worn than that of the
female. Rand (19426 : 457) suggested that the differences between these specimens
might indicate that the Dutch New Guinea birds form another race, but on this point
he was not certain. The evidence provided by the birds from the Central Highlands
suggests that these are seasonal differences. Dr. Rand also noted a slight difference
in the colour and clarity of the black markings and vermiculations of the scapulars
and inner secondaries of his birds. In all the birds of the present series these tend to
be closer to the Lake Habbema bird in colour and to the Mount Tafa bird in clarity,
so it may well be that this small sample is from a mixed population.
Collocalia esculenta esculenta (Linnaeus)
Hirundo esculenta Linnaeus, 1758, Syst. Nat. 10th ed., p. 191. " China " error for Ambon
{ex Rumphius).
Field notes : These swiftlets often occurred in pairs ; once, however, about 12
were seen hawking for insects above a native clearing on a steep hillside at about 6,000
ft.. A nest was found under a bank beside a stream, in it were two elongate eggs a
pale blue-white in colour (10th September). The nest was made of fine roots and
thickly covered with a yellowish coloured dry lichen that hung from the trees.
4 o8 BIRDS COLLECTED BY MR. F. SHAW-MAYER
Collacalia hirundinacea hirundinacea Stresemann
Collocalia fuciphaga hirundinacea Stresemann, 1914, Verh. Ornith. Ges. Bayern, 12, p. 7.
Setakwa River, Dutch New Guinea.
Field note : A few birds were seen, two being found on the nest. The nests were
built of fine rootlets, thin black stalks and a few pieces of moss with some earth
towards the bottom. An elongate egg was found in one of the nests.
Alcyone azurea ochrogaster Reichenow
Alcyone ochrogaster Reichenow, 1903, Journ. f. Ornith. 51, p. 149. Ramu River, northern New
Guinea.
1953. 17. 115 ; i ? ; Minj, Wahgi River, 5,000 ft.; November, 1950.
Measurements : 1 ? — wing, 77 ; tail, 33 ; bill, 46. Colours : bill, black ; tip
pale horn ; legs, orange-red ; iris, not recorded. Stomach contents : insects.
This bird matches ochrogaster except that the upper-parts are a duller blue like
lessonii.
Clytoceyx rex rex Sharpe.
Clytoceyx rex Sharpe, 1880, Ann. Mag. Nat. Hist. (5) 6, p. 231. East Cape, New Guinea.
1953. 17. 116 ; i ? ; Mount Wilhelm, Bismarck Range, 6,000 ft.; May, 1950.
Measurements : 1 $ — wing, 173 ; tail, 137 ; bill, 37. Colours : bill, pale horn
(base of upper mandible dark horn) ; legs, green-grey ; iris, black-brown. Stomach
contents : remains of large beetles.
This bird is more richly coloured than others I have seen of the species. This is
interesting because Paludan described septentrionalis on the paler undersurface of
three specimens from the nearby Sepik area. In size it is intermediate between rex
and imperator van Oort from the Noord River — imperator,9. : wing, 180 ; tail, 148 ;
(van Oort, 1909 : 79) — rex, $ : wing, 160-165 ; tail, 115-125.
Halcyon sancta sancta Vigors and Horsfield.
Halcyon sancta Vigors and Horsfield, 1827, Trans. Linn. Soc. London, 15 ; p. 206. New South
Wales, Australia.
1953.17.117-118 ; 2 (J ; Mount Giluwe, 7,600-7,500 ft.; May, 1951.
Measurements : 2 (J— wing, 91, 92 ; tail, 59, 60 ; bill, 35, 37. Colours : bill,
black (lower mandible pale towards base) ; legs, dark-grey ; iris, dark-brown.
These non-breeding visitors were taken in the wooded hills north of Mount Giluwe.
Both are in moult,
BIRDS COLLECTED BY MR. F. SHAW-MAYER 409
Hirundo tahitica frontalis Quoy and Gaimard
Hirundo frontalis Quoy and Gaimard, 1830, Voy. Astrolabe, Zool. 1, p. 204, pi. 12. Dorey,
north-western New Guinea.
Field Note. About 15 to 20 of these birds were to be seen on most evenings on
the earthy bank around the pond at Nondugl (8th September) .
Edolisima montona montona (Meyer)
Campephaga montona A. B. Meyer, 1874, Sitzungsber. Akad. Wiss, Wein, 69. p. 386. Arfak
Mountains, north-western New Guinea.
1953. 17. 122 ; 1 c? ; Hagen Range, 8,000 ft.; March, 1951.
Measurements : 1 <$ — wing, 135-5 \ tail, 101 ; bill, 16. Colours : bill, black ;
legs, black ; iris, black-brown. Stomach contents : berries.
Rothschild and Hartert (1907 : 464) examined birds of this species and found that
the adult males from the mountains of western New Guinea had longer wings than
those from the mountains of the south-east ; they named these shorter-winged birds
minus. This difference in wing length, however, does not appear to be so well marked
now that more material is available ; Junge (1953 : 40), for example, reported that
the wing lengths of some adult males in a series from one locality in western New
Guinea were within the range of measurements of minus. Nevertheless, until more
material is available from other localities the species cannot be reviewed satisfactorily
but it seems probable that when this is done minus will be found to be no longer a
valid race separable on wing length. The specimen from the Hagen Range is similar
in measurement to some of the birds reported by Junge.
Wing Length of Adult Males
Western New Guinea. South-eastern New Guinea.
Rothschild and Hartert . 136-5-141 -5 Rothschild and Hartert . . 128-132
Junge .... 131 -141-5 British Museum (Natural History) 125-134
Coracina longicauda longicauda (De Vis)
Grauculus longicauda De Vis, 1890, Ann. Rep. Brit. New Guinea, 1888-89, p. 59. Musgrave
Range, south-eastern New Guinea.
Field note. A nest was found about 50-60 ft. above the ground in a fork of a
horizontal branch on a somewhat isolated tree in the forest near Tomba. It was
saucer-shaped and built of fine rootlets, the outside was decorated with lichens and
moss. There was one juvenile on n October, which was fed by both parents. The
food appeared to consist mainly of winged insects, but on two occasions (13th October)
lizards were brought. After being fed with the first one, which had been pushed down
its throat with some difficulty, the young bird made a gurgling noise which may have
4 io BIRDS COLLECTED BY MR. F. SHAW-MAYER
been a begging call somewhat stifled by the full crop. On the second occasion when a
lizard was brought to the nest only the tail protruded from the adult's mouth, the
adult then regurgitated the lizard to give it to the young. When the nest was examined
on the following day (14th October) an uneaten green lizard was found there. The
nest was kept under observation for about three and a half hours and the feeding
visits that the adults paid to the nest during that time were recorded as follows :
08.35
Male and female.
10.30 :
Male.
08.37
Female.
10.42 :
Male.
08.47
Female.
11.25 :
Male and female
0943
Male and Female.
12.07 :
Male.
Nest sanitation was attempted only once, but that may have been because the
birds had been disturbed. When it was not being fed the young bird remained
wonderfully still, it never changed its position during the whole of the morning while
it was being watched from the hide. On another day the bird was moved around in
its nest so that it faced the sun, it turned immediately and resumed its former
position.
Anthus gutturalis rhododendri Mayr
Anthus gutturalis rhododendri Mayr, 1931, Mitt. Zool. Mus. Berlin, 17, p. 692. Mongi-Busu,
Saru waged Mountains (2,600 m.), north-eastern New Guinea.
1953. 17. 123-124 ; i (J, i juv. (J ; Hagen Range, 11,000 ft.; February, 1951.
1953.17.125-131 ; 4 c?, 3 ? ; Mount Giluwe, 12,000-11,500 ft.; April, 1951.
Measurements : 5 <$ l — wing, 98-101 ; tail, 74-79 ; bill, 12-14. 3 $ — wing,
94-97 ; tail, 74-77 ; bill, 13-14. Colours : bill, black ; legs, dark horn ; iris,
dark brown. Stomach contents : grass seeds and insects.
1 Measurements of 1953. 17. 124 not given owing to broken plumage.
Anthus australis exiguus Greenway
Anthus australis exiguus Greenway, 1935, Proc. New England Zool. Club, 14, p. 35. Wau,
Morobe district, north-eastern New Guinea.
1953. 17. 132-138 ; 3 <J, 3 $, 1 juv. ? ; Mount Giluwe, 7,300 ft.; May, June, 1951.
Measurements : 3 $ — wing, 84-86 ; tail, 62 ; tarsus, 26-27 ; pollux (not claw),
9-9-5 ; claw of pollux, 10-13 \ bill, 13-5-14. 3 ? — wing, 80-82 ; tail, 55-59 ; tarsus,
26 ; pollux (not claw), 9 ; claw of pollux, 11-13 ; bill, 12-14. 1 juv $ — wing, 76 ;
tail, 53 ; tarsus, 25 ; pollux (not claw), 9 ; claw of pollux, 12 ; bill, 13. Colours :
bill, dark horn (lower mandible paler towards base) ; legs, dark flesh to horn ; iris,
dark brown. Stomach contents : insects.
" This pipit was plentiful on the plateau at just over 7,000 feet. It was especially
numerous on any cleared area of grassland and in the native gardens. As many as
50 to 60 were observed in the early morning and late afternoon on a recently made
football field near lumbal." (F. S.-M.).
BIRDS COLLECTED BY MR. F. SHAW-MAYER 411
No colour difference can be seen between most of the males and females of this
series and the type and allotype respectively of exiguus kindly sent on loan to me by
Mr. Greenway, One adult male, however (1953 . 17 . 134) is slightly paler, but warmer
in colour above owing to the light brown edging to the feathers being wider than in the
type ; the breast and flanks are correspondingly warmer and are washed a light
brown. They are also slightly larger, particularly in the length of the wing and tail,
but probably not enough to be recognized racially.
Field note. During September they were seen in pairs on the lawn around the
house at Nondugl and on the grasslands where they were very common. One bird
was seen in partial display, it flew typically pipit-like into the sky and then volplaned
down.
Saxicola caprata wahgiensis Mayr and Gilliard
Saxicola caprata wahgiensis Mayr and Gilliard, 1951, Amer. Mus. Novit. No. 1524, p. 8.
Mafulu, south-eastern New Guinea.
1953. 17. 139-142 ; 2 (J, 2 $ ; Mount Giluwe, 7,300 ft.; May, June, 1951.
Measurements : 2 <J — wing, 77, 79 ; tail, 55, 56 ; bill, 11. 2 $ — wing, 77, 78 ;
tail, 56, 58 ; bill, 11. Colours : bill, black ; legs, black ; iris, brown-black.
Stomach contents : insects (grubs and adults).
" Two or three of these friendly stone-chats are usually to be seen around any
building in the grasslands on the plateau. Their song is the first bird call to be heard
in the early morning." (F. S.-M.).
Compared with the female specimen of wahgiensis collected by Mr. Gilliard at
Nondugl the two females from Mount Giluwe are slightly darker below and on the
sides of the neck. In the darker colour of the underparts these birds tend towards the
Oranje Mountain bird, belensis, of which the females are larger than the present pair,
Mayr and Gilliard (195 1 : 8).
Field note. Native name, " Kompichente ". A nest was found built on the
ground on a pile of pit-pit roots and stems. It contained two juveniles with the
feathers of the first plumage newly broken through the skin (26th September) . Only
the female was seen to visit the nest. A pair was seen in the garden perched on
sticks in the flower beds, each dropped down tc the ground from time to time to pick
up insects. A fairly tame bird often seen in the Mount Hagen area.
T urdus poliocephalus erebus Mayr and Gilliard
Turdus poliocephalus erebus Mayr and Gilliard, 1952, Amer. Mus. Novit. No. 1577, p. 7. New
name for Turdus poliocephalus carbonarius Mayr and Gilliard, 195 1, Amer. Mus. Novit. No.
1524, p. 7. Mount Orata, Kubor Mountains, Central Highlands, New Guinea. (Not
Turdus carbonarius Lichtenstein, 1823.)
1953 . 17 . 143-146 ; 3 (J, 1 $ ; Mount Giluwe, 11,000 ft.; April, 195 1.
Measurements : 3 $ — wing, 124-132 ; tail, 90-94 ; bill, 20-21. 1 ? — wing, 123 ;
tail, 92 ; bill, 21. Colours : $ — bill, orange-yellow with darker tips ; legs, orange ;
iris, brown with yellow eye ring ; inside of mouth, orange. ? — bill, upper mandible
4 i2 BIRDS COLLECTED BY MR. F. SHAW-MAYER
dark horn, lower mandible orange ; legs, dull orange ; iris, dark brown — yellow
eye ring. Stomach contents : berries and small beetles.
" On the top of Mount Giluwe the blackbird leaves the shelter of the small timber
in the late afternoon and flies into the open alpine grassland where it searches among
the rocks and tussock-grasses for its food." (F. S.-M.). The gonads of all the
specimens were enlarged.
Melampitta lugubris longicauda Mayr and Gilliard
Melampitta lugubris longicauda Mayr and Gilliard, 1952, Amer. Mus. Novit. No. 1577, p. 1.
Mount Tafa, Wharton Range, south-eastern New Guinea.
1953 . 17 . 147-148 ; i juv. (J, i ? ; Mount Giluwe, 9,000-8,500 ft.; April, 1951.
Measurements : 1 juv. <$ — wing, 83 ; tail, 33 ; tarsus, 40 ; bill, 15. 1
§ — wing, 84 ; tail, 54 ; tarsus, 39 ; bill 16. Colours : bill, black ; legs, black ;
iris, dark brown. Stomach contents : small insects.
The juvenile male is tawny black with the under-parts washed with honey-
brown. Its primaries appear to be fully grown, but the central rectrices are absent.
Crateroscelis robusta robust a (De Vis).
Gerygone robusta De Vis, 1898, Ann. Rep. Brit. New Guinea, 1896-97, p. 84. Wharton Range,
south-eastern New Guinea.
1953. 17. 149-150 \ 2 $\ Mount Giluwe, 8,000-7,500 ft.; June, 1951.
Measurements : 2 $ — wing, 64, 65 ; tail, 39, 40 ; bill, 12. Colours : bill, upper
mandible dark horn and lower mandible light horn ; legs, flesh ; iris, yellow to
orange-red.
Although these birds have been named robusta they differ from topotypical
specimens by being more greyish in colour. In addition, the wing coverts, the lower
part of the back, lower abdomen and flanks have a faint olivaceous wash. Mayr and
Gilliard (1954 : 345) and Gyldenstolpe (1955 : 84) reported that among the birds from
the Bismarck, Hagen and Kubor Mountains which they named robusta there was some
variation in colour although a few specimens matched topotypical material. This
pair from Mount Giluwe were sent to the Berlin Museum ; there Professor Stresemann
kindly compared them with the type of albigula Reichenow from Schraderberg in the
Sepik Mountains, which is now regarded as a synonym of robusta. He found that they
agreed with this specimen so it might appear that albigula is a valid race, but in view
of the variation in the series collected by Mr. Gilliard and Count Gyldenstolpe I
consider that it is better not to separate albigula on these two specimens.
Eupetes leucostictus loriae Salvadori
Eupetes loriae Salvadori, 1896, Ann. Mus. Civ. Genova, (2) 16, (= 36), p. 102. Moroka, south-
eastern New Guinea.
1953. 17. 151, 153, 153a ; 1 (J, i §," 1 juv. ^ ; Mount Giluwe, 8,500 ft.; May, June>
1951.
BIRDS COLLECTED BY MR. F. SHAW-MAYER 413
1953. 17. 152 ; 1 ? ; Hagen Range, 8,500 ft.; February, 1951.
Measurements : 1 <J — wing, 76 ; tail, 82 ; bill, 18. 2 ? — wing, 84,86 ; tail, 94,
98 ; bill, 18, 19. 1 juv. $ — wing, 82 ; tail, 84 ; bill, 17. Colours : bill, black ;
legs, brown-grey to brown-black ; iris, dark brown.
These birds are completing moult or are in fresh plumage except the February bird,
where the plumage is worn. They were taken in the forests of the north slopes of
Mount Giluwe. Nearest to loriae but differ by being a clearer blue below, due to the
green wash being restricted to the flanks, the sides of the thorax and to a small patch
on the upper breast near the throat. (The blue, however, is not so bright as that of a
specimen of amabilis received on loan from the American Museum of Natural History.)
The upper-parts appear to be a brighter green, slightly less olivaceous in colour ;
this difference, however, may be due to fading on the part of the older specimens.
The bills are somewhat more slender than in both topotypical loriae and amabilis,
particularly the latter.
Field note. Native name, " Penkeklempo " or "Kengnan" (the latter is
probably the name for E. castanonotus) . A nest was found in a hole about 2 ft. above
the ground in a half-dead stump of a tree-fern. An adult continued to visit the nest
frequently and did not appear to be shy although the porters were nearby. It
searched for insects on the ground by turning over moss and leaves with a quick
movement of the bill ; insects that were taken were too small to be identified. It is
possible that both parents feed the young because a second visit to the nest followed
too soon upon the first for the same individual to have found food so quickly. When
it returned to the nest the bird remained near the ground and jumped into the nest-
hole from immediately beneath it. Faeces were removed from the nest on several
occasions. The nest was situated so that no sunlight fell directly on it and the nest
hole faced to the east. On 5th October some of the vegetation was removed to allow
more light to fall on the nest and two young birds covered with a heavy grey down
were discovered there. They crouched down in the middle of the nest at first, then the}^
stretched out their necks and started to hiss in a snake-like manner. The nest was
then observed from a short distance away, after a while the juveniles started to preen;
but when the sun began to fall directly on to the nest the young birds began to shift
around and utter a harsh cry. Finally they jumped out of the nest and one of the
parents lead them away along a small run through the undergrowth. The nest itself
was circular in shape, with an external diameter of about 145 mm. and an internal
one of about 75 mm.. It was heavily built on an untidy foundation of sticks, ferns and
dead leaves, the inner cup was composed of fine rootlets lined with the skeletons of
leaves, although one green leaf was removed from the nest. A second nest was found
containing two young birds on 8th October ; these were about the same stage of
development as the young in the first nest. It was situated in a nest-hole about 3 ft.
from the ground and similarly built to the first one only with less foundation. Both of
the parents fed the young. The gape of the juvenile was white and the inside of the
mouth a pale orange. The young were fed eleven times in one hour at the height of
feeding, but only five times between 11.50 and 12.50 hours. On one occasion one of
the parents sat on the nest and started to brood the young, when the other came to
feed the juveniles it pushed the brooding parent off the nest ; there appeared to be
4 i4 BIRDS COLLECTED BY MR. F. SHAW-MAYER
no recognition behaviour. The adults were very tame and continued their search for
insects to within an arm's length of the hide and ignored all noises that came from
inside. A third nest was found in a hole towards the base of a tree-fern. In it were
two eggs (16th October), Mayr and Gilliard (1954 : 345).
Ifrita kowaldi kowaldi (De Vis)
Todopsis kowaldi De Vis, 1890, Ann. Rep. Brit. New Guinea, 1888-89, p. 59. Owen Stanley
Mountains, south-eastern New Guinea.
Field note. Native name, " Kompe-chicko ". A nest was found built about 12
ft. up in a slender sapling, it was thickly decorated with moss which gave it a massive
appearance. In the nest there was one juvenile which seemed to suffer badly from
the heat. Both parents fed the young bird ; once the juvenile was given a cicada-like
insect which was so large that the parent seemed to have difficulty in forcing it down
the young bird's throat.
Malurus alboscapulatus subspecies ?
Malurus alboscapulatus A. B. Meyer, 1874, Sitzungsber. Akad. Wiss. Wien, 69, p. 469. Arfak
Mountains, north-western New Guinea.
1953. 17. 154-157 ; i cJ, 2 juv. (J, i ? ; Mount Giluwe, 7,500-7,300 ft.; April, June,
1951.
Measurements : 3 <J — wing, 51-53 ; tail, 50-59 ; bill, n-5-i2-5. 1 ? — wing,
49 ; tail, 57 ; bill, 11. Colours : bill, black (unsexed specimen, lower mandible
dark horn) ; legs, dark horn ; iris, dark horn. Stomach contents : insects.
Mayr and Gilliard (1954 : 346) and Gyldenstolpe (1955 : 88) reported that apart
from wing size all the birds collected in the Wahgi region agreed with topotypical
specimens of the race mafulu from the mountains of south-eastern New Guinea. It
seems to me, however, that the adult male from Mount Giluwe is a closer match with
a specimen of the longer winged race balim from the Snow Mountains which I had on
loan from the American Musuem of Natural History than with a specimen of mafulu.
But, again, the immature birds seem to be a fairly close match with young mafulu.
The birds from Mount Giluwe are also larger than birds of the race tappenbecki (<£,
wing, 45-48) from the lower Ramu and Sepik rivers north of the Schrader-Bismarck
Range, an area considerably nearer than either the Snow Mountains or Mafulu.
Professor Stresemann who kindly compared the series with topotypical specimens of
tappenbecki in the Berlin Museum was able to separate the immature birds on colour
as well as wing length.
Megalurus timoriensis subspecies ?
Megalurus timoriensis Wallace, 1863, Proc. Zool. Soc. Lond. 1863, p. 489. Dilly, Timor.
Series A :
1953. 17. 158 ; 1 (J ; Mount Giluwe, 12,000 ft.; April, 1951.
1953 . 17 . 159-162 ; 1 juv. <J, 3 $ ; Lamende Range, 11,000 ft.; May, 1951.
BIRDS COLLECTED BY MR. F. SHAW-MAYER 415
Series B :
1953. 17. 163-166 ; 4 (J ; Mount Giluwe, 7,300 ft.; April, 1951.
Measurements : Series A : 2 <$ — wing, 73, 74 ; tail, 95-5, 96 ; bill, 12, 14.
3 ? — wing, 68-71 ; tail, 93-97 ; bill, 12. Series B : 4 <$ — wing, 71-75 ; tail,
98-109 ; bill, 12-13. Colours : bill, upper mandible dark horn and the lower
mandible pale horn ; legs, brownish dark flesh ; iris, Series A, light to dark brown —
Series B, pale orange to light brown. Stomach contents : insects.
Mayr and Gilliard (195 1 : 9) described two races of this species of warbler from
Mount Hagen. They named the birds of the summit (alpine) grasslands at 12,000 ft.
montanus and those of the grass slopes at 7,800 ft. wahgiensis. The birds were
separated mainly on the somewhat darker, more rufous, colouring of the light areas
of the upper-parts of montanus. Through the kindness of the American Museum of
Natural History I have examined two specimens of montanus and one of wahgiensis
from the series on which the descriptions were based. The specimens in Series A and
Series B listed above are distinguishable from each other and, as far as I can see, the
high altitude series matches Mayr and Gilliards' lower altitude form wahgiensis,
while the lower series on Mount Giluwe matches the high altitude montanus. The
explanation of this curious reversal on two neighbouring peaks is not evident. It
seems certain, at least, that the differences are not caused by seasonal changes in
plumage for in both series there are birds in worn and fresh dress.
Field note. Native name, " Kuchaill " (" 11 " as in Welsh).
Cisticola exilis diminuta Mathews.
Cisticola exilis diminuta Mathews, 1922, Birds Austr. 9, p. 373. Paterson Creek, Cape York,
Queensland, Australia.
1953. 17. 167 ; i ? ; Mount Giluwe, 7,300 ft.; June, 1951.
Measurements : 1 ? — wing, 44 ; tail, 47 ; tarsus, 18 ; bill, 9-5. Colours :
bill, horn colour, darker on ridge ; legs, flesh colour ; iris, dull brown. Stomach
contents : insects.
Generally regarded as being a species restricted to the grasslands below 4,000 ft.
it is interesting that this specimen was taken at 7,300 ft.. The specimen is in winter
plumage.
Sericornis perspicillatus Salvadori
Sericornis perspicillata Salvadori, 1896, Ann. Mus. Civ. Genova. (2) 16 ( = 36), p. 99. Moroka,
south-eastern New Guinea.
Field note. A somewhat pear-shaped nest was found in the forest near Tomba.
It was about 2 to 3 ft. above the ground, hanging by a few fibres from a twig. The
entrance to the nest was about half-way up one side ; above it a small bulge formed
a " porch ". Two young birds were in the nest (15th October). Both parents fed the
4 i6 BIRDS COLLECTED BY MR. F. SHAW-MAYER
young and during a period of one hour 16 visits were made with food ; nest sanitation
occurring 3 times. During another period of watching for 30 minutes food was taken
to the nest 7 times. Among the insects fed to the juveniles a moth, a small green
caterpillar and a Tipulid were identified. The adults chattered noisily around the
nest. They were active and seldom remained still, hopping around in the undergrowth
and sideways up the stems of the bushes. They flirted their wings constantly, like
birds of the genus Phylloscopus.
Gerygone ruflcollis insperata De Vis
Gerygone insperata De Vis, 1892, Ann. Rep. Brit. New Guinea, 1890-91, p. 94. Mount Suckling,
south-eastern New Guinea.
Field note. Native name, " W'un-monamp ". A domed nest containing two
fully-fledged juveniles was found on 3rd October in the forest near an area of swamp-
land. One point of interest is that three different adults appeared to feed the young
birds. Apart from this the behaviour, of the adults was typical of most warblers ;
they flitted with quick movements from branch to branch and never remained still
for any length of time. Both the young and the adults were very noisy.
Peltops montanus Stresemann
Peltops blainvillii montanus Stresemann, 1921, Anz. Ornith. Ges. Bayern, 1, p. 35. Hunstein-
spitze, Sepik Mountains, northern New Guinea.
Field note. One bird was seen in the forest at the top of a high tree, the identity
of this blackbird with a white patch below and behind each eye and the crimson
rump and vent was unmistakable.
Rhipidura brachyrhyncha devisi North
Rhipidura devisi North, 1897, Proc. Linn. Soc. New South Wales, 22, p. 444. New name for
Rhipidura albicauda De Vis, 1897, Ibis, p. 375. No type locality given (= ? Mt. Scratchley).
(Not Rhipidura albicauda North, 1895).
1953. 17. 168 ; 1 imm. <J ; Mount Giluwe, 7,500 ft.; June, 1951.
Measurements : 1 imm. <$ — wing, 71-5 ; tail, 92 ; tarsus, 24 ; bill, 8. Colours :
bill, upper mandible dark horn, lower mandible light horn ; legs, pale flesh ; iris,
brown-black. Stomach contents : insects.
Rhipidura albolimbata lorentzi van Oort
Rhipidura albo-limbata lorentzi van Oort, 1909, Nova Guinea, 9, (Zool.), p. 85. Hellwig Moun-
tains, Oranje Range, New Guinea.
1953. 17. 169 ; i ? ; Mount Giluwe, 8,000 ft.; June, 1951.
Measurements : 1 $ — wing, 82 ; tail, 85 ; bill, 10. Colours : bill, not recorded ;
legs, black ; iris, brown-black. Stomach contents : insects.
BIRDS COLLECTED BY MR. F. SHAW-MAYER 417
Field note. Native name, " Ipenem ". Two nests were found, one was built
about 20 ft. above the ground in a tree, the other about 12 ft. up in a creeper. Each
was a small neat cup with a " tail ", the outsides were close-knit in appearance as if
spiders' webs had been woven in. Both nests contained young (26th September)
which the adults continued to feed while the party stood beneath the nests.
Rhipidura leucophrys melaleuca (Quoy and Gaimard)
Muscipeta melaleuca Quoy and Gaimard, 1830, Voy. Astrolabe, Zool. 1, p. 180. New Ireland.
1953. 17. 170 ; 1 $ ; Mount Giluwe, 7,300 ft.; June, 1951.
Measurements : i $ — wing, 104 ; tail, 103 ; tarsus, 25 ; bill, 13. Colours :
bill, black ; legs, black ; iris, brown-black. Stomach contents : insects.
Usually regarded as a bird of the coast, lowland lakes, swamps and rivers it is
interesting that Mr. Shaw-Mayer found it to be "fairly plentiful" on the high
plateau near Mount Giluwe. Mayr and Gilliard (1954 : 348) reported that it was
common at Nondugl at 5,000 ft., and that it was nesting at 7,000 ft.
Field note. A common bird which spends most of the time on the ground. The
tail is not spread out much compared with many other members of the genus. When
the bird jumps up to a perch from the ground the long tail is swung pendulum-
fashion in the manner of that of a shrike.
Machaerirhynchus nigripectus saturatus Rothschild and Hartert
Machaerirhynchus nigripectus saturatus Rothschild and Hartert, 1913, Novit. Zool. 20, p. 498.
Mount Goliath, Snow Mountains, New Guinea.
1953. 17. 171 ; i c? ; Mount Giluwe, 8,000 ft.; May, 1951.
Measurements : 1 <J — wing, 64 ; tail, 62 ; bill, n. Colours : bill, black ; legs,
black ; iris, black-brown. Stomach contents : small insects.
Field note. Native name, " Koongnaremp ". Fairly common in the forest.
Microeca papanua Meyer
Microeca papuana A. B. Meyer, 1875, (April), Sitzungsber. Abh. Nat. Ges. Isis, Dresden, 1875,
p. 74. Arfak Mountains, north-western New Guinea.
I 953- I 7-i72 ; i(J; Mount Giluwe, 7,500 ft.; May, 1951.
Measurements : 1 <J- — wing, 78-5 ; tail, 48 ; bill, 10. Colours : bill, black ;
legs, yellow ; iris, dark brown. Stomach contents : insects.
This specimen is a little larger and darker in colour than the other birds of this
species that I have seen and there is marked reduction in yellow on the outer webs
of the primaries. Rand (1942& : 483) reported similar differences in the birds he
4i
BIRDS COLLECTED BY MR. F. SHAW-MAYER
collected in the mountains of Dutch New Guinea and correlated the increase in size
with altitude. He suggested that his birds appeared to be darker because the material
with which they were being compared was foxed, and this may well account for the
colour difference between the birds before me.
Petroica bivittata bivittata De Vis.
Petroeca bivittata De Vis, 1897, Ibis, p. 376. Mount Scratchley, south-eastern New Guinea.
1953. 17. 173, 176 ; 1 (J, 1 ? ; Mount Giluwe, 11,000-10,000 ft.; April, May, 1951.
1953. 17. 174-175 ; 2 (J ; Lamende Range, 11,000 ft.; May, 1951.
Measurements : 3 <J — wing, 73-77 ; tail, 46-48 ; bill, 9-0. 1 ? — wing, 73 ; tail,
45 ; bill, 9-0. Colours : bill, black ; legs, black ; iris, dark brown. Stomach
contents : small insects.
The April bird (sex ?) has started moulting, one bird taken in May is in fresh
plumage the other two are completing moult. Collected on the alpine grasslands.
Peneothello sigillatus subspecies ?
Poecilodryas (?) sigillatus De Vis, 1890, Ann. Rep. Brit. New Guinea, 1888-89, p. 59. Mount
Victoria, south-eastern New Guinea.
Field note. An adult female was seen on 18th October on an empty nest which
was half concealed in moss and creeper. The nest was built of moss, green ferns and
rootlets, with a thick lining of moss and dead fern leaves. It measured approximately
50 mm. in diameter internally and 115 mm. externally, the cup being about 25 mm.
deep.
Peneothello cyanus subcyaneus (De Vis)
Poecilodryas subcyanea De Vis, 1897, Ibis, Ser. 7, 3, p. 377. Mountains of south-eastern New
Guinea.
1953. 17. 177 ; i <J ; Mount Giluwe, 7,500 ft.; May, 1951.
Measurements : 1 $ — wing, 99 ; tail, 68 ; bill, 15. Colours : bill, black ;
legs, black ; iris, brown-black. Stomach contents : small insects.
Field note. Native name, " Goorgla ". Nests were built in the forks of upright
branches in the bushes, and appeared to be made mainly of fine rootlets covered with
moss and lined with dried fern fronds. They were round and deep, being about 55
mm. in diameter and 45 mm. deep internally and 130 mm. in diameter and 80 mm.
high externally. One nest, first examined on 18th September, contained one juvenile
almost ready to fly ; it was visited daily until 22nd September, when the young
bird had flown. Another nest had one juvenile in it on 28th September, but this bird
BIRDS COLLECTED BY MR. F. SHAW-MAYER 419
had flown when the nest was examined on the following day. An egg was found in a
nest 17th October. It was a pointed oval with a light olive ground colour ; there
Mere a few scattered dark brown spots and a fainter red-brown speckling that became
more intense towards the broader end.
Heteromyias albispecularis centralis Rand.
Heteromyias albispecularis centralis Rand, 1940, Amer. Mus. Novit. No. 1074, p. 4. Near
Bernhard Camp, Idenburg River, at 2,150 m., central New Guinea.
1953. 17. 178-180 ; i o, i juv. cJ, i $ ; Mount Giluwe, 8,000-7,800 ft.; April,
June, 1951.
Measurements : 2 <J< — wing, 99, 101 ; tail, 61, 62 ; bill, 17, 18. 1 ? — wing, 98 ;
tail, 58 ; bill, 17. Colours : bill, black with tip pale horn (juv. <J, pale horn) ;
legs, flesh ; iris, dark brown. Stomach contents : small beetles and other insects.
The two April birds are completing moult, although a few juvenile feathers still
remain in the young male. The adult male (June) is in fresh plumage.
Field note. Native name " Yamari." Several nests were found in dark parts of
the moss forest where the light-meter read as low as 3-2 foot-candles for the most of
one forenoon. They were built 7 to 10 ft. above the ground in the upper branches of
young saplings, although one was found in a small bush where it was only 3 ft. from
the ground. The nests appeared to be very similar in being heavily built on slender
branches, which in some cases could not adequately support their weight. One nest
that was examined was found to be about 90 mm. in diameter and 40 mm. deep ;
it was built of grass and twigs with a little moss and lined with thin fern rootlets. All
the nests contained young birds. One juvenile examined on 20th September was
blackish brown above with the shafts of the feathers lighter in colour. The breast
was brown and the belly white with a brown wash. The gape and the inside of the
mouth were yellow. The legs, which were long and strong, were flesh coloured.
The irides were brown. Another nest, seen two days later on 22nd September,
contained only one nestling which was quite naked ; yet a week later, on 29th
September, a nest was found with one juvenile which had well-grown wing quills.
This young bird rose on its legs and with arched wings started to cheep when the
parents returned to the nest. It was established that both parents fed the young
whose main diet appeared to consist of earthworms. On two occasions a juvenile was
seen to eject a bag of faeces on to the rim of the nest, one of the adults returned to
the nest expressly to remove it — or so it appeared. The adults were never seen in the
forest except at the nest. Sometimes they would return to the nest silently, at other
times they would call quietly with a soft, musical piping sound. The young always
answered this call, but not loudly. When the adult made this piping call the throat
vibrated, but the bill remained closed. The alarm call was, " kak-kaktak-kak ".
Once when a nest was being examined an adult repeatedly exhibited a distraction
display by " injury feigning " on the ground where it dragged one wing as if it were
broken.
zool. 3, 10. 27
420 BIRDS COLLECTED BY MR. F. SHAW-MAYER
The adults had a habit of remaining hidden in the undergrowth and of standing
quietly on the nests, a form of behaviour that is more characteristic of a babbler than
a flycatcher. The bird's long strong legs and heavy bill are other characters that
suggest that this bird may have some affinity with the Timalinae. As opposed to
these Timaloid tendencies adults were seen to perch sideways on the moss-covered
trunks of the trees in a position curiously like that of a nuthatch.
Pachycephala schlegelii obscurior Hartert
Pachycephala schlegelii obscurior Hartert, 1896, Novit. Zool. 3, p. 15. Eafa District, Owen
Stanley Mountains, south-eastern New Guinea.
1953. 17. 181 ; i (J ; Mount Giluwe, 7,500 ft.; May, 1951.
Measurements : 1 <J — wing, 88 ; tail, 65 ; bill, 11. Colours : bill, black ;
legs, brown-black ; iris, brown-black. Stomach contents : insects.
Pachycephala modest a hypoleuca Reichenow
Pachycephala hypoleuca Reichenow, 1915, Journ. f. Ornith. 63, p. 125. Schraderberg, Sepik
Mountains, northern New Guinea.
1953. 17. 182-183 ; i (J, i ? ; Lamende Range, 11,000 ft.; May, 195 1.
Measurements : 1 <J — wing, 90 ; tail, 64 ; bill, n. 1 ? — wing, 89 ; tail, 65 ;
bill, 12. Colours: bill, black; legs, dark grey to black ; iris, dark brown. Stomach
contents : small grubs and insects.
These specimens were collected among the small trees on the alpine grasslands.
Pachycephala rufinucha niveifrons Hartert
Pachycephala rufinucha niveifrons Hartert, 1930, Novit. Zool. 36, p. 57. Wondiwoi, Wandam-
men Mountains, north-western New Guinea.
1953.17. 184 ; 1 juv. <J ; Hagen Range, 8,000 ft.; March, 1951.
1953. 17. 185-186 ; 2 (J ; Mount Giluwe, 8,000-7,8000 ft.; April, 1951.
Measurements : 3 <J — wing, 89-91 ; tail, 68-70 ; bill, 16-5, (juv., 14). Colours :
bill, black (juv., horn) ; legs, olivaceous grey-black (juv., horn) ; iris, yellow (juv.,
dark grey). Stomach contents : insects.
Apart from having slightly longer wings these birds match material taken at lower
altitudes on the Utakwa River (wing, 84-86). Rand (19426 : 489) showed that in this
species wing length could be correlated with altitude, the birds from the higher
altitudes being slightly larger.
Field note. A nest was built in a fork of a branch about 7 ft. up in a bush. The
outer part of the nest was made of ferns, coarse rootlets and green moss, the inner
part of the thin fibres, fine rootlets and the skeletons of leaves. There were two well
BIRDS COLLECTED BY MR. F. SHAW-MAYER
421
incubated eggs in the nest on 17th October. These were a chalky white colour with
black spots and a fainter underlay of grey spots. Another nest found two days later,
on 19th October, contained two juveniles ; the insides of their mouths were a pale
orange apart from two prominent blue spots at the posterior of the palate. It is
probable that the young were fed by both parents, but the nest was visited only
infrequently. During a three and a half hour watch on the nest an adult visited it
only four times during the first half hour, three times during the second, once during
the third but four times during the last half-an-hour.
Myiolestes megarhynchus tappenbecki (Reichenow)
Colluricincla tappenbecki Reichenow, 1899, Journ. f. Ornith. 47, p. 118. Friedrich Wilhelms
Hafen (= Madang), Astrolabe Bay, north-eastern New Guinea.
1953. 17. 187-190 ; 2 cJ, 2 ? ; Minj, Wahgi River, 5,000 ft.; November, 1950.
1953. 17. 191 ; 1 ? ; Mount Giluwe, 7,5000 ft.; May, 1951.
Measurements : 2 & — wing, 94, 96 ; tail, 76, 79 ; bill, 20, 21. 3 ? — wing, 91 —
92 ; tail, 72 — 76 ; bill, 18 — 19. Colours : bill, 2 £, brown horn — 3 $, horn ; legs,
blue-grey to dark grey ; iris, brown. Stomach contents : insects ; 1 $, small
(?) water snail.
All the four birds taken at Minj in November, 1950, were taken on the nest, the
nests being built about one foot above the ground in long grass. It is interesting to
note that these four birds are in different plumages. The two males are in full moult ;
and of the two November females one is in worn dress while the other is in fresh
plumage. The plumage of the female taken in May shows some signs of wear.
Eulacestoma nigropectus subspecies ?
Eulacestoma nigropectus De Vis, 1894, Ann. Rep. Brit. New Guinea, 1893-94, p. 102. Mount
Maneao, south-eastern New Guinea.
1953. 17. 192 ; ? ? ; Mount Giluwe, 8,000 ft.; May, 1951.
Measurements : 1 ? ? — wing, 71 ; tail, 52 ; bill, 11-5. Colours : bill, horn ;
legs, purple-flesh ; iris, grey-brown. Stomach contents : insects.
This is a juvenile, and in the absence of comparable material I cannot name it
racially. It is probably clara Stresemann and Paludan as Mayr and Gilliard (1954 :
354) reported that their birds from this area were that race.
Lanius schach stresemanni Mertens
Lanius schach stresemanni Mertens, 1923, Senckenbergiana, 5, p. 228. Kulungtufu, Saruwaged
Mountains, north-eastern New Guinea.
1953. 17. 193, 195-196 ; 1 <J, 2 ? ; Mount Giluwe, 7,300 ft.; May, June, 1951.
1953. 17. 194 ; 1 juv. cJ ; Wahgi River, 5,200 ft.; November, 1950.
ZOOL. 3, IO . 27§
422 BIRDS COLLECTED BY MR. F. SHAW-MAYER
Measurements : i $ — wing, 101 ; tail, 140 ; bill, 16. 1 juv. <J — wing, 95 ;
tail, 105 ; bill, 13. 2 $ — wing, 97, 98 ; tail, 121, 123 ; bill, 15. Colours : bill,
black (juv. (J, lower mandible, horn) ; legs, black ; iris, brown-black. Stomach
contents : small grubs, beetles and other insects.
" Not many of this solitary species on the grasslands of the plateau. A bird here
and there, as usual clinging to the stem of cane-grass." (F. S.-M.). The adult male
is in fresh plumage and the testes are slightly enlarged. The plumage of the young
male is worn. The two females are completing moult with the outer primaries
growing.
Field note. Native name, " Komkompir ". Two deep-cupped nests, built about
6 ft. above the ground, were found in tall reeds. One, examined on 5th September,
contained two eggs; but in the other, found on 23rd September in pit-pit country,
were two well-grown juveniles.
Artamus maximus Meyer
Artamus maximus A. B. Meyer, 1874, Sitzungsber. Akad. Wiss. Wien, 69, p. 203. Hatam,
Arfak Mountains, north-western New Guinea.
1953. 17. 197 ; i c? ; Minj, Wahgi River, 5,000 ft.; November, 1950.
Measurements : 1 (J- — wing, 164 ; tail, 73 ; bill, 23. Colours : bill, blue-grey ;
legs, grey ; iris, dark brown. Stomach contents : small beetles.
Gyldenstolpe (1955 : 121) separated A. m. wahgiensis mainly on the darker colour
of the upper-parts of the birds from the Wahgi region in comparison with a series from
the Arfak Mountains. The present specimen, however, hardly differs from the Arfak
Mountain bird. The only variations in colour in the specimens I have examined
appear to be correlated with age and season. In a series from south-eastern New
Guinea there are some individuals lighter in colour than both the Wahgi River and
Arfak birds, and others that are darker in colour. It seems that the lighter coloured
birds are juveniles or adults in worn, faded plumage ; while the darker birds are
adults in fresh plumage. Unfortunately this series is incomplete in specimens taken
in all months of the year, so the degree of seasonal variation cannot be established
with certainty. As the individual variation is greater than that between the birds
from the Central Highlands and the Arfak Mountains, I am not naming the present
bird racially.
Field note. Native name, " Kompurkaikir ". A nest was built about 50 ft.
above the ground in the fork at the base of an upright branch of a dead tree in the
forest. Three juveniles, almost ready to fly, were found in it (26th September). It is
interesting that four or five different adults were observed to feed the young, and on
one occasion three adults were seen on the nest at the same time ! Communal nesting
of this nature is rare and provides a subject for further investigation. Like many
other communities this one was very noisy, with both the adults and the juveniles
contributing to the chatter. The juvenile plumage was similar to that of the adult,
but the dark areas of the body were necked with a light brown ; the bill of the young
bird was horn coloured.
BIRDS COLLECTED BY MR. F. SHAW-MAYER 423
Aplonis metallica metallica (Temminck)
Lamprotornis metallicus Temminck, 1824, Planch. Col. d'Ois, pi. 266. " Celebes," by error for
Amboina, Moluccas.
1953. 17.198 ; i ? ; Mount Giluwe, 7,000 ft.; May, 1951.
Measurements : 1 ? — wing, 104 ; tail, 91 ; bill, 16. Colours : bill, black ;
legs, black ; iris, bright orange-red.
" Few Aplonis were seen at 7,000 feet when this specimen was taken in the wood-
lands but the bird was plentiful at 5,300 feet." (F. S.-M.).
Pomareopsis briujni (Salvadori)
Grallina bruijni Salvadori, 1875, Ann. Mus. Civ. Genova, 7, p. 929. Arfak Mountains, north-
western New Guinea.
Field note. Native name, " Notechll " (" 11 " as in Welsh). This is a bird of the
mountain stream, and one was seen by a rushing torrent at about 6,500 ft. When it
alighted on a rock it fanned out its tail in the same manner as the Plumbeous Red-
start, Pheonicurus fuliginosus , of the fast-flowing streams of the Himalayas.
Epimachus meyeri bloodi Mayr and Gilliard
Epimachus meyeri bloodi Mayr and Gilliard, 1951, Amer. Mus. Novit. No. 1524, p. 10. Mount
Hagen, Central Highlands, 8,300 ft., New Guinea.
1953. 17. 199-200 ; i (J, i juv. (J ; Hagen Range, 9,000 ft.; February, 1951.
1953. 17.201 ; 1 ? ; Mount Wilhelm, Bismarck Range, 8,000 ft.; May, 1951.
Measurements : 1 # — wing, 174 ; tail, 604 ; bill, 78. 1 juv.<J — wing, 149 ;
tail, 320 ; bill, 60. 1 °- — wing, 149 ; tail, 312 ; bill, 76. Colours : bill, black ; legs,
dark grey (ad. <$ black) ; iris, blue-grey. Stomach contents : fruits.
The juvenile male resembles a series of adult females from the Hagen Range except
in the length of the bill. The adult female from the Bismarck Range differs from this
series in the upper-parts being lighter, less olivaceous, more like the ochraceous
colour of the females of meyeri. Not until more material is available from the Bis-
marck Range will it be possible to assess the importance of this difference, but it may
be that the Wahgi Gap country plays just as an important part in the distribution of
the races of E. meyeri as it does in the distribution of the races of Astrapia stephaniae,
(see below). The plumage of the present birds is fairly fresh, showing little signs of
wear. Specimens taken in October in previous collections from Mount Hagen are in
worn plumage and those taken in November are in full moult. It appears from the
stomach contents of the birds that E. meyeri is not entirely frugivorous, for the
stomachs of adult birds have been found to contain berries, grasshoppers and other
insects in addition to various fruits.
424 BIRDS COLLECTED BY MR. F. SHAW-MAYER
Astrapia stephaniae femina Neumann
Astrapia stephaniae femina Neumann, 1922, Verh. Ornith. Ges. Bayern, 15, p. 236. Schrader-
berg, Sepik Mountains, 60 miles north of the Hagen Mountains, northern New Guinea.
1953.17.202 ; 1 $ ; Mount Wilhelm, 8,000 ft.; May, 1950.
Measurements : 1 ? — wing, 145 ; tail, 320 ; bill, 24. Colours : bill, black ;
legs, blue-grey (feet, black) ; iris, black-brown. Stomach contents : fruits.
Mayr and Gilliard (1952a) showed that the grasslands between the Hagen Range
and the mountains of the Wahgi Divide (a westerly spur of the Bismarck Range)
acted as a barrier in the distribution of A. mayeri, a species closely related to A.
stephaniae. The same authors (1954 : 321) also described the deforestation in the
Wahgi Valley, which they suggested formed another important barrier to forest
birds. It appears, then, that the mountain forest birds of the Kubor Range along the
south and the Hagen Range at the head of the Wahgi Valley are isolated by open
country from the mountain forest birds of the Wahgi Divide and the Bismarck Range
north of the valley. This barrier apparently influences the distribution of the races
of A. stephaniae, for specimens collected in the Bismarck Range by Mr. F. Shaw-
Mayer in 1946 have been matched with cotypical material of femina, and Gylden-
stolpe (1955 : 127) identified his birds from north of the Wahgi Valley as this race.
On the other hand, two specimens from Mount Giluwe (see below) appear to be
ducalis, which is the race found to the south of the valley in the Kubor Range (Mayr
and Gilliard, 1954 : 356). The adult females from the Bismarck Range are slightly
more olivaceous above than the adult females of ducalis and the under-parts are rust
coloured barred with black instead of a buffy colour barred with black. In the colour
of the under-parts they agree with the differential characters given by Mayr and
Gilliard (1954 : 356). (They do not, however, exhibit the subobsolete white streaks
at the bases of the central tail feathers described by these authors.) The present
specimen differs somewhat from other material in having a narrow chestnut collar
barred with black across the shoulders, the colour of the collar becoming lighter
laterally as it merges into the rust barred-black of the breast.
Astrapia stephaniae ducalis Mayr
Astrapia stephaniae ducalis Mayr, 1931, Mitt. Zool. Mus. Berlin, 17, p. 711. Dawong, Herzog
Mountains, eastern New Guinea.
1953.17.203-204 ; i (J, i $ ; Mount Giluwe, 8,500 ft.; March, April, 1951.
Measurements : 1 $ — wing, 165 ; tail, 660 ; bill, 20. 1 $ — wing, 137 ; tail,
320 ; bill, 21. Colours : bill, black ; legs, dark grey ; iris, brown-black. Stomach
contents : fruits and berries.
Field note. Five birds, females or immature males, were seen in the forest. In
flight, with their long tails streaming behind, they were reminiscent of the Racquet-
tailed Drongo, Dissemurus paradiseus ; the diving flight being particularly like that
BIRDS COLLECTED BY MR. F. SHAW-MAYER 425
of the Dicruridae. The birds appeared to be hunting for insects in the moss covering
the branches of the trees ; one bird was seen to swallow what was probably a spider.
Astrapia mayeri Stonor
Astrapia mayeri Stonor, 1939, Bull. Brit. Ornith. Club, 59, p. 57. " Eighty to a hundred miles
west of Mount Hagen," central New Guinea.
1953.17.205 ; 1 c? ; Mount Giluwe, 9,000 ft.; March, 1951.
1953.17.206-207 ; 1 juv. (J, 1 ? ; Hagen Range, 9,000-8,500 ft.; December, 1950.
Measurements : 1 cJ — wing, 177 ; tail, 852 ; bill, 13. 1 juv. <$ — wing, 170 ; tail,
350 ; bill, 12. 1 ? — wing, 148 ; tail, 310 ; bill, 15. Colours : bill, black ; legs,
blue-grey (feet, black) ; iris, brown-black. Stomach contents : fruits and green
berries.
Field note. The branches of the forest trees on which the bird perches are
thickly covered with moss. It is probably this layer of moss that prevents the long
tail feathers from becoming damaged.
Astrapia stephaniae x Astrapia mayeri
1953.17.208 ; 1 (J ; Hagen Range, 8,000 ft.; March, 1951.
Measurements : 1 $ — wing, 167 ; tail, 821 ; bill, 18. Colours : bill, black ;
legs, dark grey (feet black) ; iris, brown-black.
Close to mayeri in having the tail white for most of its length ; but the rachis of
the distal two-thirds of the central rectrices are brown in colour, and the webs of
these feathers are fringed irregularly with brown. The nasal tuft is reduced in size
to that of an immature male of mayeri, that is intermediate between the condition in
meyeri where the tuft is present and in stephaniae where it is absent. The primaries
are darker than in mayeri, being a solid black with a purple lustre like those of
stephaniae. Too much importance should not be attached to the proportion of hybrid
specimens in collections, for in the field they occur in a limited area which happens
to be one of the most accessible. Mr. Shaw-Mayer wrote (in correspondence 5th
November, 1952) of the difficulties he had experienced in obtaining a series of hybrids,
and that, " Over the greater part of their extensive range the species remain distinct.
The overlapping areas where hybridism takes place are small. ... ".
Paradisaea apoda salvadorii Mayr and Rand
Paradisaea apoda salvadorii Mayr and Rand, 1935, Amer. Mus. Novit. No. 814, p. 11. Vanumai,
Central Division Papua, south-eastern New Guinea.
1953.17.209. i $ Wahgi River, 5,500 ft.; November, 1950.
Measurements : 1 ? — wing, 161 ; tail, 118 ; bill, 32. Colours : bill, blue-grey ;
legs, brown-horn ; iris, bright yellow. Stomach contents : fruits.
426 BIRDS COLLECTED BY MR. F. SHAW-MAYER
" Count Raggi's Bird-of-Paradise was fairly plentiful on the Wahgi River plateau ;
finding sanctuary in the casuarina groves, small stands of secondary forest and in the
few remnants left of the original oak forests. By mid-November (1950) they were
breeding and several nests were observed ; in nearly every case the nests were high
up, 50 to 60 feet, in casuarina trees. Each nest contained only one egg." (F. S.-M.).
Pteridophora alberti hallstromi Mayr and Gilliard
Pteridophora alberti hallstromi Mayr and Gilliard, 1951, Amer. Mus. Novit. No. 1524, p. 12.
Near Tomba, Hagen Range, Central Highlands, New Guinea.
1953. 17. 210 ; i juv. (J ; Mount Wilhelm, 7,500 ft.; May, 1950.
1953. 17. 211 ; 1 ? ; Hagen Range, 9,000 ft.; February, 1951.
Measurements : 1 juv. <? — wing, 112 ; tail, 80 ; bill, 15. 1 $ — wing, 122 ; tail,
81 ; bill, 15. Colours : bill, black ; legs, brown-black (juv. $, grey-brown) iris,
dark brown ; inside of mouth of $, green. Stomach contents : green berries.
Field note. Native name, " Kongbuk " (adult male); " Doongbi " (adult
female and juveniles). An adult male was seen in the forest at about 7,000 ft. The
bird was preening and called occasionally, the sound was like the squeaking of rusty
iron. The long head plumes were manoevured to avoid the branches of the tree. A
solitary bird.
Loria loriae amethystina Stresemann
Loria loriae amethystina Stresemann, 1934, Ornith. Monatsber. 42, p. 144. Schraderburg, Sepik
Mountains, northern New Guinea.
1953. 17.212 ; 1 (?; Mount Wilhelm, 6,500 ft.; May, 1950.
1953. 17. 213 ; 1 cJ ; Hagen Range, 9,000 ft.; February, 1951.
Measurements : 2 <J -wing, 103, 106 ; tail, 71, 72 ; bill, 15-5, 16. Colours :
bill, black ; mouth- wattles, creamy white ; legs, dark olive ; iris, dark brown.
Stomach contents : berries.
Field note. A domed nest with an exceptionally wide round opening was built
about 12 ft. up in a decayed tree-stump in the forest. It was deep-set in living moss
and built of green ferns and moss with a lining of fern stalks. There was one juvenile
in the nest (25th September). Its head and breast were a smoke-grey and the wings a
darker grey ; the gape and the inside of the mouth were white in colour. While the
nest was being watched during the early part of one morning the female returned
about every 10 to 15 minutes to feed the young and called with a soft " wark, wark "
each time she approached. The food was regurgitated and appeared to consist of
large berries and plum-coloured fruits. A short while after being fed the young bird
expelled the fruit stones from its crop, they were ejected with a force sufficient to
shoot them out through the mouth of the nest. Faecal capsules were also ejected
occasionally from the nest, but generally the female assisted with nest sanitation.
When the nest was examined a few days later it was deserted and contained faeces and
fruit stones.
BIRDS COLLECTED BY MR. F. SHAW-MAYER 427
Cnemophilus macgregorii kuboriensis Mayr and Gilliard
Cnemophilus macgregorii kuboriensis Mayr and Gilliard, 1954, Bull. Amer. Mus. Nat. Hist. 103,
p. 361. Mount Orata, Kubor Mountains, Central Highlands, New Guinea, + 9,000 ft.
1953. 17.214 ; 1 (J ; Minj River, Kubor Range ; 8,000-9,000 ft.; October, 1950.
Measurements : i $ — wing, 109 ; tail, 88 ; bill, 13 ; crest, 37. Colours : bill,
black ; legs, dark olive ; iris, dark brown. Stomach contents : fruits.
Mayr and Gilliard separated this race on the paler colour of the upper-parts in
comparison with sanguineus from Mount Hagen. In colour it is closer to the scarlet-
vermilion of sanguineus than the pale orange of macgregorii from south-eastern New
Guinea ; the colour of kuboriensis being orange- vermilion. This specimen was taken
near the headwaters of the Minj River and it matches a series from the Bismarck
Range, which suggests that there may be an east to west cline from the pale colour of
macgregorii in the south-east to the deep, rich colour of sanguineus from the Hagen
Range. The series consists of birds in new and worn plumage but there is little
seasonal colour difference.
Cnemophilus macgregorii sanguineus Iredale
Cnemophilus macgregorii sanguineus Iredale, 1948, Australian Zool. 11, p. 162. Kumdi, Mount
Hagen district, Central Highlands, New Guinea.
1953. 17.215 ; i juv. <J ; Hagen Range, 9,000 ft.; February, 1951.
1953. 17.216 ; 1 imm. <J ; Mount Giluwe, 9,000 ft.; May, 1951.
Measurements : 1 imm. $ — wing, 109 ; tail, 85 ; bill, 20 ; crest, 36. 1 juv.
cJ- — wing, 113 ; tail, 87 ; bill, 20 ; crest, 21. Colours : bill, black (juv. (J, tip
dark horn) ; legs, brown-black to glossy black ; iris, dark brown (juv. <J, dull grey).
Stomach contents : fruits.
The young bird is a juvenile and indistinguishable from an adult female. The
immature bird could well be described as " sub-adult ", for in colour it is intermediate
between the young bird and an adult male, but the collector has indicated on the
field label that the testes were as large as those of a breeding adult male. From this it
seems that the bird comes into breeding condition before it is fully adult. If this is
so it would be interesting to determine its behaviour towards an adult female and
her response. The primaries and rectrices of this specimen are as richly coloured as
topotypical sanguineus ; that is, darker than kuboriensis where these feathers are
nearer the colour of macgregorii. It is on this character that I have been able to
determine the racial identity of the bird.
Archboldia papuensis sanfordi Mayr and Gilliard
Archboldia papuensis sanfordi Mayr and Gilliard, 1950, Amer. Mus. Novit., No. 1473, p. 1. Four
miles west of Tomba, south-western slope of Mount Hagen, Central Highlands, New Guinea.
1953. 17.217, 218, 220, 222; 3 <J, 1 imm. <J ; Mount Giluwe, 9,000-8,500 ft.;
March, April, May, 195 1.
428 BIRDS COLLECTED BY MR. F. SHAW-MAYER
1953. 17. 219, 221, 223, ; i (J, i juv. (J, i imm. $ ; Hagen Range, 9,000-8,800 feet ;
April, 1951.
Measurements : 4 <J — wing, 165-171 ; tail, 167-171 ; bill, 17-20. 1 juv.
(J — wing, 142 ; tail, 131 ; bill, 19. 1 imm. $ — wing, 163 ; tail, 141 ; bill, 22. 1 imm.
$ — wing, 154 ; tail, 132 ; bill, 19. Colours : bill, black ; legs, grey-blue (imm. ?,
bright light blue) ; iris, dark brown. Stomach contents : fruits.
These are the first specimens of this genus to be received by the British Museum
(Natural History). In comparison with the description of this race the feathers of
the crest in the male exhibit some variation. Although they are predominantly
black-based with orange-gold tips some have an area of white between the black and
the orange-gold and a few are tipped with black. All the specimens are in moult.
Field note.- — Near Tomba an area of the forest floor had been cleared to form a
bower measuring approximately 5 ft. by 4 ft. The branches of the surrounding trees
and a fallen tree trunk were densely decorated with the bamboo-like trailing vines of
epiphytes which grow on high trees. Several branches were quite bare and smooth,
probably from wear when the birds alighted and postured. The floor of the bower
was strewn with the dead fronds of ferns. A number of shells of land snails were also
found on the ground.
Amblyornis macgregoriae De Vis
(Text-fig 2)
Amblyornis macgregoriae De Vis, 1890 (22nd February), Ann. Rep. Brit. New Guinea, 1888-89,
p. 61. Musgrave Range, south-eastern New Guinea.
1953.17.224 ; 1 cJ ; Rubor Range, 7,000 ft.; November, 1950.
1953.17.225-226 ; i<$, 1 juv.cJ; Hagen Range, 9,000 ft.; February, 1951.
1953.17.227 1 (J ; Lamende Range, 9,000 ft.; June, 1951.
Measurements : 4 <J — wing, 134-137 ; tail, 83-86 ; bill, 17-19 ; crest, 60-65.
Colours : bill, dark horn lighter towards base ; legs, dark olive-brown ; iris, dark
brown. Stomach contents : fruits.
Mayr and Gilliard (1954 : 363) discussed the small variations in colour and measure-
ment found in this species. They considered that these were of a clinal nature and
concluded that because of this it would be inadvisable to distinguish their birds from
the Central Highlands as a separate race, although they differed somewhat from other
populations. The material I have examined supports this conclusion, the underparts
of the birds from the Central Highlands being, on the whole, rather more ashy in
tone when compared with other populations. This difference however, is not constant
and there is some individual variation for two adult males from Tomba match birds
from the south-east. Although I have few adult males for comparison their measure-
ments do not differ significantly from those of the birds from the Central Highlands.
Measurements of adult males of Amblyornis macgregoriae
Wing. Tail. Crest.
macgregoriae Owen Stanley Range (2) . . 132, 140 . 80 . 57, 61
aedificans Upper Waria River (2) , , 132, 138 . 80, 81 . 60, 61
BIRDS COLLECTED BY MR. F. SHAW-MAYER
429
^•>&-jv>V/:.>: : :-:;y:i : :-::^<:i\^
2^i 260 .4-»--230 — *c
,1050
Fig. 2. — Sketch of a cross-section of the bower of Amblyornis macgregoriae.
Measurements in millimetres. (After Loke Wan Tho.)
Field note. Bowers were found on several coccasions ; they were roughly
circular in plan, but the dancing ground invariably formed a perfect circle. Each
bower was built around a central sapling of Garcenia which the natives call ' ' Kichan' ' .
One bower found deep in the moss forest was built of a dark moss and the playground
was devoid of leaves.
Chlamydera lauterhachi lauterbachi Reichenow
Chlamydera lauterbachi Reichenow, 1897, Ornith. Monatsber.
Ramu River, Central Highlands, New Guinea.
14. Jagei River, upper
1953.17.228-230 ; i (J, 2 ? ; Minj, Wahgi River, 5,000 ft.; November, 195 1.
Measurements : 1 £ — wing, 133 ; tail, 104 ; bill, 22. 2 ? — wing, 126, 129 ;
tail, 102, 103-5 J bill, 22. Colours : bill, black ; legs, olive-green ; iris, brown.
Stomach contents : fruits and berries.
" This bower-bird is found in lightly wooded country. It was plentiful on the
plateau drained by the Wahgi River, where it frequented small patches of secondary
forest and the shrub around the edges of fields. It was often seen flying low from one
small group of trees to another. The birds were breeding in November and several
nests were observed. The normal clutch size appeared to be one egg. The bower is
roughly circular in shape and about two feet in diameter. Inside there are two main
runways approximately parallel to each other with their openings converging on
opposite sides of the bower, The walls are about one foot high and built of interlaced
43Q BIRDS COLLECTED BY MR. F. SHAW-MAYER
sticks. The runways, or ramps, incline upwards from the openings towards the middle
of the bower where, at their highest, they are joined by a smaller transverse runway ;
thus in plan they form an H. The transverse runway is about ten inches long with
walls about eight inches high built of very small sticks and twigs. A small heap of
blue-coloured stones was found in this transverse runway and more stones and a few
dark blue fruits in the main ones." (F. S.-M.). The design and construction of the
bower agrees with the description given by Marshall (1954).
Daphoenositta miranda kuboriensis Mayr and Gilliard
Daphoenositta miranda kuboriensis Mayr and Gilliard, 1952, Amer. Mus. Novit. No. 1577, p. 5
Mount O-Mar, Kubor Mountains, Central Highlands, New Guinea.
1953. 17.231-233 ; i <J, i (?) imm. (J, 1 ? ; Lamende Range, 10,000 ft.; May, 1951
Measurements : 1 (J — wing, 87 ; tail, 43 ; bill, 10-5. 1 (?) imm. <J — wing, 81
tail, 42 ; bill, 9-0. 1 ? — wing, 82 ; tail, 40 ; bill, io-o. Colours : bill, black
legs, black (?, yellow) ; iris, cream ((?) imm. (J, yellow). Stomach contents : small
insects.
From the description of this race it seems that the doubtfully sexed specimen may
be a female as the feet and legs are yellow in colour, whereas they are dark green
in an adult male. Its plumage is similar to that of other adults, so it is unlikely to
be a young bird. The plumage is fairly fresh in all the birds and there is no sign of
moult. The specimens were taken in the wet moss forest high on the Lamende Range.
Neositta papuensis alba Rand
Neositta papuensis alba Rand, 1940, Amer. Mus. Novit. No. 1072, p. 10. 15 km. south-west
of Bernhard Camp, Idenburg, River, northern New Guinea.
Neositta papuensis wahgiensis Gyldenstolpe, 1955, Ark. f. Zool. 8, p. 153. Nondugl, Wahgi
Valley, Central Highlands, New Guinea.
1953.17.234-236 1 cJ, 2 juv. <J ; Mount Giluwe, 8,000 ft.; June 1951.
Measurements: i $ — wing, 84; tail, 38; bill (from nostril), io-o. 2 juv.
cJ — wing, 76-5, 78-5 ; tail, 34, 40 ; bill (from nostril), 9, 11. Colours : bill, pale
yellow with black tip ; legs, yellow ; iris, yellow. Stomach contents : insects.
The adult male matches a topotypical male taken by Dr. Rand. It is characteristic
of alba in having the head and neck pure white all round and the feathers of the
breast and abdomen paler edged than in papuensis. The female of alba is unknown
but Gyldenstolpe (1955 : 153) described a single female from Nondugl which he
distinguished as wahgiensis. It seems probable that this female may represent the
hitherto unknown female of alba because of the identity of the males from Mount
Giluwe ; but this point cannot be settled until topotypical females of alba have been
collected. Meanwhile, I propose that wahgiensis should be regarded as a synonym of
alba.
BIRDS COLLECTED BY MR. F. SHAW-MAYER 431
Field note. Three birds were seen in a tree where they appeared to be hunting
for insects. They worked their way up and along the branches searching carefully
and at long length among the moss.
Myzomela adolphinae Salvadori
Myzotnela adolphinae Salvadori, 1875, Ann. Mus. Civ. Genova, 7, p. 946. Arfak Mountains
north-western New Guinea.
Field note. A solitary bird. Two or three would usually be seen at intervals
along the drive of the bungalow where we stayed at Nondugl. Each bird would perch
on the top of one of the small trees of the casuarinas hedge.
Myzomela rosenbergii wahgiensis Gyldenstolpe
Myzomela rosenbergii wahgiensis Gyldenstolpe, 1955, Ark, f. Zool. 8, p. 155. Weiga, Sepik-
Wahgi Divide, Central Highlands, New Guinea (7,400 ft.).
Field note. Native name " Kaltentepal ". A fairly common bird often seen
flying from tree to tree in small flocks of a dozen or so, and heard calling out most of
the time. A nest was found about 10 ft. above the ground in a small bush growing in
a swamp, it was built of roots and lined with finer rootlets. Two young birds were
found there (24th September), these were a dark chestnut brown above and a lighter
brown below with a pale brown patch under the chin. The wings were a dark chestnut
brown with a darker barring and the flight feathers black in colour. The gape and
the inside of the mouth were yellow and the eyes brown, the legs were grey in colour,
but the backs of the legs and the feet were yellow. Only the female was observed to
feed the young birds, no male was seen to go to the nest. The juveniles defaecated
with considerable force so that the faeces were ejected clear of the nest.
Melipotes fumigatus Meyer
Melipotes fumigatus A. B. Meyer, 1886, Zeitschr. ges. Ornith. 3, p. 22. Hufeisenbirge, south-east-
ern New Guinea.
1953.17.237-238 ; i ^, i $ ; Hagen Range, 7,000 ft.; December, 1950 ; January,
1951.
Measurements : 1 <J — wing, 124 ; tail, 108 ; bill, 21. 1 $• — wing, 115 ; tail,
100 ; bill, 20. Colours : bill, black ; legs, grey-blue ; iris, dark brown ; skin of
face, orange. Stomach contents : berries and fruits.
The specimens match the description of the large-sized birds reported from the
same locality by Mayr and Gilliard (1954 : 366) who also described a smaller and
somewhat darker bird from the neighbouring Wahgi-Divide. They did not distinguish
432 BIRDS COLLECTED BY MR. F. SHAW-MAYER
the birds trinominally because they were of the opinion that the differences were
due not to geographical but to local altitudinal variations. In not naming these
specimens racially I am following their decision. Gyldenstolpe (1955 : 157) identified
the smaller bird as goliathi Rothschild and Hartert.
Field note. Native name, " Mongkulle ". Several nests were found slung from
the forks at the ends of the branches of the trees. They were between 15 and 20 ft.
above the ground and made of leaves lined with rootlets and covered on the outside
with moss. One nest contained an egg on 23rd September which had a beige-pink
ground colour and red-brown spots that were more numerous towards the broader
end. Another nest contained one juvenile on 7th October ; this was fed by only one
of the parents, the other remained nearby and called unceasingly with a plaintive
squeaking, " Wheet-wheet-wheet ". The food given to the young appeared to be
fruit, on two occasions plum-coloured fruit was clearly seen to be offered. The species
was fairly common, the flight being somewhat similar to that of a bulbul. It is
probable that the patch of bare skin around the eye changes colour. (Mayr and
Gilliard, 1954 : 367, reported a change in colour of the skin of this area after a bird
had hung upside down for a short time. — R. W. S.)
Melidectes fuscus fuscus (De Vis)
Acanthochoera fusca De Vis, 1897, Ibis, Ser. 7, 3, p. 383.- Mount Scratchley, south-eastern New
Guinea.
1953.17.239 ; i (J ; Hagen Range, 10,000 ft.; February, 1951.
1953.17.240-242 ; 1 (J, 1 juv. (J, 1 ? ; Mount Giluwe, 11,000-10,000 ft., April,
May, 1951.
I 953- I 7-243 ; 1 ? ; Lamende Range, 10,000 ft., May, 1951.
Measurements : 2 <J- — wing, 108, no ; tail, 109, no ; total culmen, 29 ; tarsus,
33, 34. 1 juv. (J — wing, 96 ; tail, 99 ; total culmen, 27 ; tarsus, 29. 2 ? — wing,
95, 99 ; tail, 96, 100 ; total culmen, 28 ; tarsus, 31, 33. Colours : bill, black ;
legs — 2 <J, pale blue-grey- — 1 juv. <£, dark horn — 2 ?, dark grey ; iris, dark brown ;
skin posterior to eye, brick red ; eye-wattle, pale blue (in juv. $ both skin posterior
to eye and the eye-wattle are pale yellow) . Stomach contents : insects.
Mayr and Gilliard (1954 : 367) showed that the birds of this species in the Wahgi
region are intermediate between the nominate form from south-eastern New Guinea
and occidentalis from the Oranje and Nassau Ranges, the latter being separated on
its smaller size. One female (May) taken on a nest containing a young bird is in
worn plumage. The February and April birds are in moult, while the only May male
is in fresh plumage.
Field note. Native name, " Orbooi ". A nest containing one juvenile was found
in the forest on 8th October. It was built of fine rootlets and lined with the red fibres
of the frond of the tree-fern, the outside was decorated with moss. The juvenile was
black in colour except for a blue-grey area of bare skin around the eye and other
patches of bare skin near the ear and on the neck. The bill was black, the gape yellow
BIRDS COLLECTED BY MR. F. SHAW-MAYER 433
and the tongue was brush-like — the same as the tongue of the adult bird. The legs
were a blue-flesh colour. Both parents fed the young bird, unlike the adults of M.
belfordi and Melipotes fumigatus, where only one of the parents did so. Unlike the
former they were very tame and the adults continued to feed the juvenile while the
observer was nearby. The food that was presented to the young bird consisted of
insects and probably some nectar. The female was recognizable ; her wattles were
very small and barely perceptible.
Melidectes princeps Mayr and Gilliard
Melidectes princeps Mayr and Gilliard, 1951, Amer. Mus. Novit. No. 1524, p. 13. Mount Wilhelm,
Bismarck Mountains, Central highlands, New Guinea.
1953.17.244 ; ? ; Minj River, Kubor Range, 9,000 ft.; October, 1950.
1953.17.245 ; ? ; Hagen Range, 10,000 ft.; January, 1951.
Measurements : 2 ?■ — wing, 112, 113 ; tail, 105, 106 ; bill, 42, 44. Colours :
bill, black ; legs, pale grey to blue ; iris, dark brown ; skin posterior to eye, bright
yellow ; wattle cream.
These are probably adult females. They appear to be too small to be males (adult
male : wing, 1 17-123 mm.) and are probably not young birds because the bearded
parts of the chin and throat are white in colour, whereas this area is a yellowish-white
in immature birds — Mayr and Gilliard (1954 : 13).
Melidectes belfordi (De Vis)
Melirrhophetes belfordi De Vis, 1890, Ann. Rep. Brit. New Guinea, 1888-89, p. 60. Mount
Knutsford, south-eastern New Guinea.
1953.17.246-248; 3 (J; Hagen Range, 7,500 ft.; December, 1950; January,
1953.17.249-250 ; 2 cJ ; Mount Giluwe, 7,800-7,500 ft.; April, 1951.
Measurements: 3 $ — (Nos. 1953.17.246, 247, 250) wing, 144-153; tail, 120-
126 ; bill, 40-42. 2 $ — (Nos. 1953.17.248, 249) wing, 138, 139 ; tail, no, 112 ; bill,
40, 42. Colours : 3 $ (Nos. 1953.17.246, 247, 250) — bill, black ; legs, olive ; iris,
dark brown ; skin around eye, pale blue. 2 $ (Nos. 1953.17.248, 249) — bill, pale
blue-grey ; legs, pale blue-grey ; iris, dark brown ; gape-wattle, cream ; throat-
wattle, brick red ; skin around eye, pale green-blue. Stomach contents : berries,
small insects and spiders.
The series consists of birds which differ markedly in appearance ; namely, " black-
bills " {belfordi) and " wattle-birds " (ritfocrissalis) . These differences are so distinct
that in the past belfordi and rufocrissalis were placed in different genera — Melidectes
and Melirrhophetes respectively — and until recently were regarded as members of a
super-species ; however, Mayr and Gilliard (1952c) concluded that they were con-
specific. The two forms are allopatric and generally the " black-bills " replace the
" wattle-birds " above a critical altitude of 2,000 to 2,500 metres, where hybrids arc
434
BIRDS COLLECTED BY MR. F. SHAW-MAYER
found grading between the two phenotypes. The present series was taken near the
critical hybridization altitude and each bird exhibits some degree of heterogeneity.
By adopting the scoring system used by Mayr and Gilliard (1952c) to indicate the
degree of hybridization it is possible to show how much the individual specimens
differ from the phenotypes. Assuming that a true " black-bill " scores o and a true
" wattle-bird " 100, the birds before me score as follows : —
1953.17.246 .
• 3
1953.17.247 .
. n
1953.17.250 .
. 22
1953.17.248 .
. 60
1953.17.249 .
. . 89
Generally Nos. 246, 247, 250 could be regarded as " black-bills " and the other
two specimens, Nos. 248 and 249, as " wattle-birds ".
Field note. Native name, " Arglua ". Several nests were found built from 8
to 15 ft. above the ground in the slender forks towards the end of the branches. They
looked like woven baskets of fine grasses and fern stalks, covered and suspended by
moss. One nest, first examined on 4th October, was kept under observation ; it
contained only one juvenile. The young bird had a black head with a fringe of sandy
coloured down around the crown and a lighter coloured patch around each eye. The
upper-parts were dark grey and the under-parts somewhat lighter in colour, with
white streaks on each side of the chin ; the under-tail coverts were cinnamon. The
bill was mainly grey in colour with some white on it. Both of the parents fed the
young bird, but one did so less frequently than the other and usually remained close
by the nest. The food appeared to consist mainly of insects ; but it is probable the
nectar was given also because sometimes one of the adults would return to the nest
and stand over the juvenile and rapidly flick its long reddish tongue in and out of
the young bird's mouth. When neither of the adults could be seen they could still be
heard nearby making clucking noises ; generally they were very noisy. One loud call
that they made frequently was one of the common calls of the forest. Their alarm
call could not be established with certainty, but it appeared to be either, " Quick-
quick-quick-quick, kiow-kiow-kiow-kiow ' ' , or, " tolle-tolle-tolle, kiow-kiow-kiow-
kiow " ; in response to this call the young bird would freeze in the middle of the nest
and close its eyes. They were very shy birds and easily disturbed.
Oreornis subfrenatus salvadorii (Hartert)
Ptilotis salvadori Hartert, 1896, Novit, Zool. 3, p. 531. Owen Stanley Mountains, south-eastern
New Guinea.
1953. 17.251 ; i c? ; Mount Giluwe, 7,800 ft.; April, 1951.
Measurements : 1 <J — wing, 105 ; tail, 95 ; bill, 21. Colours : bill, black (gape,
yellow) ; legs, blackish yellow ; iris, dark brown ; bare skin behind eye, bright
yellow. Stomach contents : small insects.
BIRDS COLLECTED BY MR. F. SHAW-MAYER 435
Nearest to salvadorii but slightly lighter in colour ; it differs from melanolaema in
being darker — the under-parts being less brownish and more greyish olive in colour.
I am indebted to Professor Stresemann, who kindly compared the bird with material
of the latter in the Berlin Museum.
Ptiloprora guisei umbrosa Mayr
Ptiloprova guisei umbrosa Mayr, 1931, Mitt. Zool. Mus. Berlin, 17, p. 666. Schraderberg (2,000
ft.), Sepik Mountains, Central Highlands, New Guinea.
1953.17.252-253 \ 2 $ \ Mount Giluwe, 7,500 ft.; June, 1951.
Measurements : 2 ad. $ — wing, %%, 91 ; tail, 76 ; bill, 21, 22. Colours : bill,
black ; legs, grey-blue ; iris, latteine green. Stomach contents : small insects.
Field note. Native name, " Tongkla ". A nest was found suspended on a living
vine of Hoya. It was built of rootlets and dried leaves of the climbing bamboo and
lined with the hairs from a tree-fern. The outside of the nest was decorated with
living moss apart from one part that was covered by a broad leaf of the vine. The
adult often held its tail high like a pipit as it jumped from branch to branch, some-
times it would dive to the ground in pursuit of insects and carry them to the nest.
There were young in the nest on 10th October and both parents shared in rearing
them ; the parents fed the young by clinging upside down to the vine. The call was
a clear whistle, " Choo-weei-00 ", but as they approached the nest the adults made
a "chucking" sound.
Paramythia montium montium De Vis
Paramythia montium De Vis, 1892, Ann. Rep. Brit. New Guinea, 1890-91, p. 95. Mount
Suckling, south-eastern New Guinea.
1953.17.267, 270-271 ; 1 $, 2 ? ; Mount Giluwe, 9,500-9,000 ft.; March, 1951.
1953.17.268-269 ; 1 juv. <J, 1 $ ; Hagen Range, 9,500 ft.; January, February,
1951.
Measurements : 1 $ — wing, 106 ; tail, 100 ; bill, 16. 1 juv. $ — wing, 102 ; tail,
95 ; bill, 15. 3 ? — wing, 105-106 ; tail, 89-98 ; bill, 15-16. Colours : bill,
black ; legs, black ; feet, black. Stomach contents : fruits and berries.
Field note. Native name, " Kai-ingkun ".
Zosterops novaeguineae wahgiensis Mayr and Gilliard
Zosterops novaeguineae wahgiensis Mayr and Gilliard, 1951, Amer. Mus. Novit. No. 1524, p. 14.
Nondugl, Wahgi Valley, Central Highlands.
1953.17.254 ; 1 (J ; Wahgi River ; 5,200 ft.; November, 1951.
Measurements : 1 $ — wing, 62-5 ; tail, 42-5 ; bill, n. Colours : bill, dark
horn paler at base ; legs, grey ; iris, yellow-brown. Stomach contents : berries.
Field note. A flock of about 15 to 20 of these small birds was seen near the camp
at Tomba.
436 BIRDS COLLECTED BY MR. SHAW-BAKER
Erythrura trichroa sigillifera (De Vis)
Lobospingus sigillifer De Vis, 1897, Ibis, Ser. 7, 3, p. 389. Mountains of south-eastern New
Guinea, apparently Mount Scratchley.
1953.17.255, 257; 1 (J, 1 juv. $; Hagen Range, 7,500 ft.; December, 1950;
January, 1951.
1953.17.256 1 (J Mount Giluwe 8,500 ft.; May, 1951.
Measurements : 2 <$ — wing, 60, 61 ; tail, 44 ; bill, 12. 1 juv. ? — wing, 58 ; tail,
39 ; bill, 11. Colours : bill, black ; legs, pale horn ; iris, brown-black. Stomach
contents : small berries and seeds.
Lonchura spectabilis wahgiensis Mayr and Gilliard
Lonchura spectabilis wahgiensis Mayr and Gilliard, 1952, Amer. Mus. Novit. No. 1577, p. 7
Kegalsugl, Mount Wilhelm, Bismarck Mountains, Central Highlands, New Guinea; 8,000 ft.
1953.17.258 ; i ? ; Minj, Wahgi River, 5,000 ft.; November, 1950.
Measurements : 1 $■ — wing, 49 ; tail, 32 ; bill, 10-5. Colours : bill, blue-grey ;
legs, dark blue-grey ; iris, black. Stomach contents : grass seeds.
Seen in large flocks on the grasslands. This specimen was taken on a nest containing
four eggs.
Oreostruthus fuliginosus hagenensis Mayr and Gilliard
Oreostruthus fuliginosus hagenensis Mayr and Gilliard, 1954, Bull. Amer. Mus. Nat. Hist. 103,
p. 372. Mount Hagen, Central Highlands, New Guinea.
1953.17.259-265 ; 4 (J, 3 imm. $ ; Mount Giluwe, 11,500-9,000 ft.; April, May,
June, 195 1.
Measurements : 4 £ — wing, 67-68 ; tail, 44 ; bill, 9-5-10. 3 imm. <$ — wing,
66-68 ; tail, 40-44 ; bill, 8-5-9-5. Colours : 4 $ — bill, coral red ; legs, brown-ho^n ;
iris, orange-red. 3 imm. <J — bill, black (gape, yellow) ; legs, brown-horn ; iris,
brown. Stomach contents : grass seeds.
This race was separated on two specimens from Mount Hagen which differed,
among other characters, from fuliginosus De Vis and pallidus Rand in the back
being thinly diffused with blood-red markings. Now more material is available it
seems that this character is not constant, for only one adult male in this series has
any red on the back and then it is seen as a wash which is restricted to a small area.
In other respects these birds match the description of hagenensis. The immature
birds differ from the adults in the under-parts not being marked with blood-red. In
comparison with fuliginosus the immature birds are darker above and lighter below.
The adults are completing moult and the testes are enlarged. This fine series was
taken on the mid-mountain and alpine grasslands of Mount Giluwe.
BIRDS COLLECTED BY MR. SHAW-BAKER 437
REFERENCES
Amadon, D. 1943. Birds collected during the Whitney South Sea Expedition, LIL Amer.
Mus. Novit. No. 1237 : 1-22.
Greenway, J. C. 1935. Birds from the coastal range between the Markham and the Waria
Rivers, north-eastern New Guinea. Proc. New England Zool. Club, 14 : 15-106.
Gyldenstolpe, N. 1955- Notes on a collection of birds made in the Western Highlands,
Central New Guinea, 1951. Ark. f. Zool., 8 : 1-181.
Hartert, E. 1930. List of birds collected by Ernst Mayr. Novit. Zool. 36 : 27-128.
Iredale, T. 1948. A Check List of the Birds of Paradise and Bower Birds. Australian Zool.
2 : 163-189.
Junge, G. C. A. 1937. The birds of South New Guinea. Part I, Nonpasseres. Nova Guinea,
New Ser., 1 : 126-184.
1939- The birds of South New Guinea. Part II, Passeres. Nova Guinea, New Ser.,
3 : 1-94.
1952. New subspecies of birds from New Guinea. Zool. Meded. Leiden, 31 : 247-250.
1953- Zoological results of the Dutch New Guinea Expedition, 1939. No. 5 Zool.
Verhandl. Leiden, 20 : 1-77.
Laurie, E. M. O. 1952. Mammals collected by Mr. Shaw-Mayer in New Guinea, 1932-1949.
Bull. Brit. Mus. {Nat. Hist.), Zool. 1 : 271-318.
Marshall, A. J. 1954. Bower Birds, Biol. Rev. 29 : 1-45.
Mathews, G. M. 1916. Birds Austr. (Witherby, London), 5 : 1-440.
Mayr, E. 1931. Die Vogel des Saruwaged- und Herzoggebirges (N. O. Neu Guinea). Mitt.
Zool. Mus. Berlin, 17 : 639-723.)
1938. Birds collected during the Whitney South Sea Expedition, XL. Amer. Mus. Novit.
No. 1007: i-n.
1940. Birds collected during the Whitney South Sea Expedition, XLI. Amer. Mus.
Novit. No. 1056 : 1-12.
1941a. Birds collected during the Whitney South Sea Expedition, XLV. Amer. Miis.
Novit. No. 1133 : 1-8.
19416. List of New Guinea Birds. (The American Museum of Natural History). New
York.
Mayr, E. and Gilliard, E. T. 1950. A new Bower Bird (Archboldia) from Mount Hagen,
New Guinea. Amer. Mus. Novit. No. 1473 : 1-3.
195 1. New species and subspecies of birds from the highlands of New Guinea. Amer.
Mus. Novit. No. 1524 : 1-15.
1952a. The Ribbon-tailed Bird of Paradise (Astrapia mayeri) and its allies. Amer. Mus.
Novit. No. 1551 : 1-13.
1 952ft. Six new subspecies of birds from the highlands of New Guinea. Amer. Mus.
Novit. No. 1577 : 1-7.
1952c. Altitudinal hybridisation in New Guinea Honeyeaters. Condor, 54 : 325-337.
1954- Birds of Central New Guinea. Results of the American Museum of Natural History
Expeditions to New Guinea in 1950 and 1952. Bull. Amer. Mus. Nat. Hist. 103 : 317-374.
Mayr, E. and Rand, A. L. 1936. Results of the Archbold Expeditions, No. 10. Amer. Mus.
Novit. No. 868 : 1-3.
1937- Results of the Archbold Expeditions, No. 14. Bull. Amer. Mus. Nat. Hist. 73 :
1-248.
Ogilvie-Grant, W. R. 1915. Report on the birds collected by the British Ornithologists'
Union Expedition and the Wollaston Expedition in Dutch New Guinea. Ibis, Jubilee
Suppl., 1915 : 1-236.
Oort, E. D. van. 1909. Birds from south-western and southern New Guinea. Nova Guinea,
Zool. 9 : 51-107.
438 BIRDS COLLECTED BY MR. F. SHAW-MAYER
Rand, A. L. 1936. Results of the Archbold Expeditions, No. 12. Amer. Mus. Novit. No.
890 : 1-14.
1940a Results of the Archbold Expeditions, No. 25. Amer. Mus. Novit. No. 1072 : 1-14.
19406. Results of the Archbold Expeditions, No. 27. Amer. Mus. Novit. No. 1074 : 1-5.
1941a. Results of the Archbold Expeditions, No. 32. Amer. Mus. Novit. No. 1102 : 1-15.
19416. Results of the Archbold Expeditions, No. 33. Amer. Mus. Novit. No. 1122 : 1-2.
1942a. Results of the Archbold Expeditions, No. 42. Bull. Amer. Mus. Nat. Hist. 79 :
289-366.
19426. Results of the Archbold Expeditions, No. 43. Bull. Amer. Mus. Nat. Hist. 79 :
425-516.
Rothschild and Hartert, E. 1907. Notes on Papuan birds. Novit. Zool. 14 : 433-483.
Sims, R. W. 1954. New race of Button-quail (Turnix maculosa) from New Guinea. Bull.
Brit. Orn. CI. 74 : 37-40.
2 8 APR 1956
PLATE 13.
Eupetes leucostictus. Adult, nest and nestlings. Wahgi region, Central Highlands, New
Guinea, 8,000 ft. October 5th, 1952. Photo : Loke Wan Tho.
Bull. B.M. (N.H.) Zool. 3, 10.
PLATE 13.
EUPETES LEUCOSTICTUS.
PLATE 14.
Melidectes fuscus. Adult, nest and nestling. Tomba, Hagen Range, Central Highlands,
"New Guinea ; 8,000 it. October 9th, 1952. Photo : Loke Wan Tho.
Bull. B.M. {N.H.) Zool. 3, 10
PLATE 14
MELIDECTES FUSCUS.
2^i
2 6 apr'wse
INDEX TO VOL. Ill
The page numbers of the principal references and the new taxonomic names are printed in
Clarendon type.
Abralia
. 264
apus, Lepidurus
4.50
Abraliopsis
261, 264
apus, Lepidurus apus
5i
abyssi, Leuckartiara
• 347
archboldi, Eurostopodus
. 407
abyssi, Perigonimus
■ 338, 347
archboldi, Lycornis
• 407
Acanthachoera
• 432
Archboldia
. 427
Accipiter
• 394
archeri, Ophioglyeera
67, 107
accraensis, Amaea .
68, 139
Architeuthidae
. 266
Acholoe astericola
66, 78
Architeuthis .
259,
261, 266,
276, 278
adolphinae, Myzomela
. 431
arcticus, Lepidurus
4, 8, 52
adusta, Po villa
• 3 IQ
arctica, Yoldia
• 346
Aegotheles
406
Ardetta
• 393
Aepypodius .
. 396
arenosus, Stylarioides
68, 128
africana, Phyllodoce
66, 85
arfaki, Strix tenebricosa
• 405
africana, Sternaspis scuta
ita . . 68, 134
arfaki, Tyto tenebricosa
. 405
Aglaophamus
67, 102
arfakianus, Aepypodius
. 396
Agonus
• 343
arfakianus, Talegallus
• 396
alalunga, Germo
263, 272
argo, Argonauta
260, 275,
276, 279
alba, Neositta papuensis
391, 430
Argonauta
260, 275,
276, 279
alberti, Vitreledonella
272
Argonautidae
• 275
albicauda, Rhipidura
416
Armandia
68, 130
albifrons, Lumbrineris
67, 119
arnoldi, Hydroides
69, 144
alboscapulatus, Malurus
. 414
Artamus
422
alcocki, Sabellaria spinul(
:>sa . . 68, 134
Astatoreochromis .
298,
299, 325
Alcyone
391, 408
Astatotilapia
312, 314
aldrovandi, Eledone
. 289
Astrapia
39i,
423. 424
Alepisaurus
259-278
Astur .
• 394
allmani, Hydractinia
351, 352-36i
Atractylis
338-347
Alloposidae .
. 274
Audouinia
67, 125
Alloposus
262, 274
australiensis, Triops
4,48
alluaudi, Astatoreochrom
is . . 298, 325
australiensis, Triops australiensis
49
alta, Aphrodite
65, 73
australis, Robsonella
. 163
Amaea .
68, 139
Amblyornis
428, 429
americanum, Poly prion
. 268
Bagrus . .
• 310
amethystina, Loria loriae
. 426
banksi, Onychoteuthis . 261, 266,
276, 279
Amphinema .
338-345
bartrami, Ommastrephes
. 268
Anas
39i, 394
batesoni, Lepidurus
54
annectens, Cyrtocara
• 305
Bayonia
299-311
annectens, Haplochromis
• 305
beccarii, Chalcophaps
401
antarcticum, Rhizorhagh
im . . 347
beccarii, Gallicolumba beccarii
. 401
antarcticus, Perigonimus
. 338, 347
Bela ....
359. 360
antennata, Eunice .
67, 111
belfordi, Melidectes
. 433
Anthus
410
belfordi, Melirrhophetes
• 433
Aphanopus .
259, 267, 269
Berlesiana
192
Aphrodite
65, 73
bernhardus, Eupagurus .
• 343
apicatus, Perigonimus
338, 347
Bhawania
66,81
Aplonis .
• 391, 423
bicolor, Haplochromis
299. 3
01,
3". 32i
Apomatus
69, 144
bicolor, Macropleurodus .
2
98,
301-332
Aprasia
• 365-38 5
bicolor, Panthalis .
65, 74
44°
INDEX
bicolor, Paratilapia
. 301
Chrysococcyx
. 404
bilobatus, Lepidurus
53
cidaritis, Perigonimus
339, 345, 346
Bimeria ....
339, 345
cidaritis, Thamnostoma
• 346
bipennata, Pterolysippe
68, 136
ciliatum, Baccinum
• 358
bitentaculata, Atractylis
338, 347
cinereus, Lepidochiton
• 345
bitentaculatus, Perigonimus
. 338
Circus ....
• 395
bivittata, Petroica bivittata
39i, 418
cirrata, Berlesiana .
• 193
bloodi, Epimachus meyeri
. 423
Cirratulus
67, 127
boa, Sthenelais
65, 76
cirratus, Halitholus
337» 339, 347
Bolitaenidae
271
cirratus, Perigonimus
• 339, 347
bonelliana, Histioteuthis 259, 2
61, 267, 276
cirrhosa, Eledone
283-291
borealis, Podocoryne
• 35i
Cisticola
• 391, 415
Bougainvillia
337-347
claparedii, Leocrates
66, 87
Bougainvillidae
• 346
Clarias ....
• 310
bougainvillii, Cyanaea
• 34i
clathratus, Trophon
• 359
Bougainvillinae
• 346
clathrus, Trophon
• 358, 360
Brachioteuthidae .
. 267
Clytoceyx
.408
Brachioteuthis
261, 267
Cnemophilus .
. 427
britannica, Bougainvillia
• 34i
Cnestrostoma
3i9, 321
briujni, Pomareopsis
. 423
coccinea, Atractylis
• 347
bruijni, Grallina .
• 423
coccinea, Eunice
67, 112
brunnea, Opisthosyllis
66, 90
collaribranchis.Phyllamphicteis
68, 135
Buccinum ....
358, 360
collaris, Lysidice
67, 114
burgersi, Psittacella
• 403
Collocalia
407, 408
Colluricincla
. 421
Cacomantis ....
. 404
complanata, Eurythoe .
66, 83
calliaris, Gadus .
. 268
conferta, Dicoryne
• 339, 347
Campanopsinae .
• 347
confertum, Eudendrium .
• 347
campbelli, Polypus
• 163
confertus, Perigonimus .
. 339
Campephaga . . . .
. 409
conger, Conger
• 273
cancriformis, Triops
4. 6, 41
convoluta, Glycera
67, 104
cancriformis, Triops cancriformis
43
Coracina
. 409
canestrinii, Epicrius . . ]
71, 183-185
cornuta, Syllis
66, 90
captus, Rallus pectoralis
. 398
coronata, Chromidina
. 291
carbo, Aphanopus . 259, 2
67, 269, 273
Corsystes
• 345
carbonarius, Turdus poliocephalus .
. 411
Coryne
• 35i
caribaea, Onychia
261, 265
Cranchia
262, 270
caribaea, Teleoteuthis
265
Cranchiidae .
270
carica, Hydractinia . . 3
51, 355-36i
crassilabris, Haplochromis
3 ! 6-33°
carnea, Podocoryne
35i, 354
Crateroscelis
. 412
caroli, Ommastrephes
. 268
Cucioteuthis
261, 265
casamancensis, Potamilla
68, 141
curtus, Sipho
• 355
Casaurius ....
39i, 393
cuvieri, Octopus
260, 273
cassivelaunus, Corsystes .
• 345
Cyanaea
• 34i
castanonotus, Eupetes .
• 4i3
cyclura, Leachia
270, 277-279
Casuarius ....
• 393
cymoctypus, Phasmatopsis
260, 271
Catablema ....
• 338
cymoctypus, Taonius
. 271, 278
cataphractus, Agonus
• 343
Cyrtocara
• 305
cavernarum, Eugamasus
191-192
Cytaeidae
• 346
centralis, Heteromyias albispecularis
. 419
Cytaeis
344, 345
Chalcites ....
• 404
Chalcophaps .
. 401
Daphoenositta . . .
. 430
Charadrius ....
. 401
dartevellei, Marphysa
67, 113
Charmosyna ....
402
Dasychone
68, 140
chilotes, Haplochromis .
• 329
decorans, Perigonimus .
340, 348
chinensis, Excalfactoria .
. 397
degeni, Platytaeniodus .
398-315-332
Chiroteuthis
261, 269
Delma
• 369
Chlamydera ....
• 429
Delphinus
263, 268
Chloeia ....
66, 81
delphis, Delphinus
263, 268
Chromidina
. 291
denticulata, Berlesiana
193-196
INDEX
441
devisi, Rhipidura brachyrhyncha
diaphana, Bolitaenella
diaphana, Eledonella
diaphana, Japetella
Dicoryne
Dicyemennea
. 416
. 271
272
261, 262, 271, 277-279
• 339, 347
291
didymobranchiata, Hermodice carunculata 66, 83
diminuta, Cisticola exilis
Dinema
dinema, Amphinema
dinema Stomotoca
Diopatra
dispar, Heteroteuthis
Doratopsis
dubia, Scoloplos
ducalis, Astrapia stephaniae
Dupetor . . .
dussumieri, Tetronychoteuthis
echinata, Hydractinia
Edolisima
edwardi, Porzana tabuensis
ehlersiaeformis, Pionosyllis
Eledone
eledones, Dicyemennea
Eledonella
Enoploteuthis
Epicriidae
Epicrius
Epimachus .
erebus, Turdus poliocephalus
eremita, Onuphis
ernsti, Falco peregrinus .
Erythrura
esculenta,Collocalia esculenta
esculenta, Hirundo
Eteone
Eudendrium
Eulacestoma
Eunice .
Eupagurus
Eupetes
eupomatoides, Sabellaria
Eurostopodus
Eurythoe
Eusigalion
Euthalenessa
Excalfactoria
excelsa, Psittacella picta
excitus, Cacomantis pyrrhophanus
exiguus, Anthus australis
Exogene
exophthalmica, Doratopsis
Falco ....
falsa, Nereis
femina, Astrapia stephaniae
ferox, Alepisaurus
filamentosa, Eunice
filiformis, Cirratulus
391, 415
• 34°
34 2 . 344
• 340. 347
67, 115
261, 262, 277, 279
269
67, 123
. 424
• 39i, 393
259, 261, 276, 278
354, 355, 359
• 4°9
. 399
66, 92
283-291
291
262, 272
261, 265
171-200
174-200
• 423
. 411
67, 118
391, 395
. 436
. 407
• 407
66, 85
339-347
. 421
67, 110
• 343
412, 413
68, 133
• 407
66, 83
65,75
65,74
• 397
. 403
. 404
. 410
66, 92
269
39i, 395
66, 93
39i, 424
259-278
67, 112
67, 127
filigera, Audouinia
67, 126
filliouxii, Sepia
260, 262
flavida, Bougainvillia
• 34i
flavida, Perigonimus
• 337
fontaniana, Robsonella .
163
forbesi, Loligo
260, 264, 276
forbesi, Rallicula .
. 400
formosum, Rhizorhagium
• 347
formosus, Perigonimus .
340, 347
frontalis, Hirundo tahitica
. 409
fuliginosus, Pheonicurus .
• 423
fulva, Pluvialis dominca
. 401
fulvus, Charadrius .
401
fumigatus, Melipotes
. 431
fusca, Acanthochoera
• 432
fuscus, Melidectes fuscus
. 432
fusiformis, Owenia
68, 132
Gabbia
. 310
Gadus ....
. 268
gallapagensis, Pseudonereis
66, 99
Gallicolumba
401
Garcenia
. 429
geayi, Pareulepis .
66, 79
gelatinosus, Perigonimus
340, 347
gemmifera, Exogene
66, 92
geometricus, Epicrius
. 171
georginae, Perigonimus .
• 340, 347
georginae, Stomotoca
• 340. 347
Germo
• 263
Gerygone
412, 416
giluwensis, Turnix maculosa
• 39i, 397
glaciale, Baccinum
• 358
glauerti, Aprasia striolata
369, 378
Glycera
67, 103
goliathina, Charmosyna papou
. 402
goliathina, Charmosyna stellae
402
Goniada
67, 105
Goniadopsis .
67, 106
goodei, Bhawania .
66, 81
goreensis, Harmothoe
66, 80
gouldi, Ardetta
• 393
gouldi, Dupetor rlavicollis
^91, 393
gracilis, Eunice
67, 112
gracilis, Syllis
66, 89
Grallina
• 423
Grampus
. 268
granarius, Triops .
4,44
grandis, Oreopsittacus arfaki .
. 402
Graneledone
• 285
granulatus, Octopus
i5 2 , 158
Grauculus .
. 409
grimaldi, Lepidoteuthis .
259, 261, 276, 278
griseus, Grampus .
. . 268
grubei, Pista
68, 137
hagenensis, Oreostruthus fuligii
iosus 436
Halcyon . . . .
. 408
Haleciidae .
• 347
442
INDEX
Halecium .....
• 343
lamonti, Synoicus ypsilophorus
. 396
Halitholus .....
337-347
Lamprotornis ....
• 423
Halla
67, 120
Lanius ......
. 421
hallstromi, Psittacella modesta
. 404
Laonome .....
68, 139
hallstromi, Pteridophora alberti
. 426
Lar ......
• 339
Haplochromis
298-332
lauterbachi, Chlamydera
• 429
Harmothoe .
66, 80
lauterbachi, Chlamydera lauterbachi
. 429
Hemichromis
319, 321
Leachia .....
270
Hemitilapia .
299, 301
Leanira .....
65.77
Hermione
65,73
Leocrates .....
66, 87
Hermodice
66, 83
Lepidochiton ....
• 345
Heteromyias
• 419
Lepidonotus .....
66, 80
Heteroteuthis
261, 262
Lepidoteuthis ....
259, 261
hirundinacea, Collocalia fuciphaga .
. 408
Lepidurus .....
■ 3-54
hirundinacea, Collocalia hirundinacea
. 408
leptura, Enoploteuthis . 261, 265
277, 278
Hirundo ....
407, 409
Leuckartiara ....
337-347
Histioteuthidae
. 267
Lialis ......
• 369
Histioteuthis
259
261, 267
lignarius, Scaphander
34°» 34i
hombergii, Nephthys
67, 102
limicola, Sthenelais
65,76
Hoplotilapia .
298
-319-333
linearis, Atractylis
34L 346
hupferi, Lepidonotus
66, 80
linearis, Bougainvillia
• 346
huttoni, Robsonella
. 165
linearis, Perigonimus
337, 341
hyalina, Syllis
66, 90
linguistica, Orbinia foetida
67, 122
Hydractinia .
35i-36i
Lobospingus .....
• 436
Hydrissa
• 361
Loimia .....
68, 138
Hydroides
69, 143
Loliginidae .....
• 263
hydrophanum, Buccinum
• 360
Loligo ..... 260
261, 263
hypoleuca, Pachycephala modesta .
. 420
Loligopsis .....
260
Hypsagonus ....
• 343
Lonchura .....
• 436
hystrix, Hermione ....
65,73
longicauda, Coracina longicauda
. 409
longicauda, Grauculus
• 409
Ifrita ......
414
longicauda, Melampitta lugubris
. 412
impatiens, Lumbrineris .
67, 118
longicaudatus, Triops
• 4,46
incerta, Goniadopsis
67, 106
longicaudatus, Triops longicaudatus
47
incrustans, Salmacina
69, 142
lorentzi, Rhipidura albolimbata
. 416
infernalis, Vampyroteuthis
• 277
Loria ....
. 426
inflatus, Perigonimus
340, 347
loriae, Eupetes
. 412
insignis, Aegotheles insignis
. 406
loriae, Eupetes leucostictus
. 412
insignis, Euthalenessa
65,75
lubbocki, Lepidurus apus
5i
insperata, Gerygone ruficollis .
. 416
lucullana, Dasychone
68, 140
intermedia, Armandia
68, 130
Lumbrineris
67, 118
intermedius, Triops longicaudatus
47
Lycornis
. 407
intoshi, Sabellaria spinulosa
68, 134
Lymnorea
• 344
ishmaeli, Haplochromis .
• 325
lynchi, Lepidurus .
53
Isolda ......
68, 136
lyrochaetus, Aglaophamus
Lysidice
67, 102
67, 114
Japetella . . . . 261,
262, 271
japonica, Cytaeis ....
• 344
macgregoriae, Amblyornis
428, 429
japonica, Leanira ....
65, 77
Machaerirhynchus
. 417
jattai, Teleoteuthis
• 265
macloviana, Bougainvillia
34 1 , 346
jeanneli, Astatotilapia . . 314
3i5>3i8
maclovianus, Perigonimus
341, 346
jeanneli, Haplochromis
• 3i5
Macropleurodus . . . 298
299-332
jonesii Perigonimus
340, 347
macrops, Haplochromis . . 308,
3H, 3i5
macropus, Octopus
273, 279
kowaldi, Ifrita kowaldi
. 414
maculosa, Turnix ....
• 397
kowaldi, Todopsis
• 4H
madagascarensis, Scoloplos
67, 123
kuboriensis, Cnemophilus macgregorii
. 427
major, Neopsittacus musschenbroekii
. 402
kuboriensis, Daphoenosit
ta miranda
. 430
Maldane ....
68, 131
Malurus
414
INDEX
443
margaritifera, Pyroteuthis
Marphysa
Mastigoteuthis
materfamilias, Haplochromis .
materfamilias, Hemitilapia
mauretanicus, Triops cancriformis
maximus, Artamus
mayeri, Astrapia . . .
mayri, Porzana pusilla .
medusa, Loimia
Megalurus ....
melaleuca, Muscipeta
melaleuca, Rhipidura leucophrys
Melampitta ....
melanocephala, Per
Melidectes
Melipotes
Melirrhophetes
Melosira
menzeli, Epicrius
metallica, Aplonis metallic a
metallicus, Lamprotornis
meyeri, Astrapia
meyeri, Epimachus
meyerii, Chalcites .
meyerii, Chrysococcyx
michaeli, Haplochromis .
Microeca
miniata, Atractylis
miniatus, Perigonimus
minor, Epicrius
minuta, Hydractinia
minutus, Perigonimus
mollis, Alloposus .
mollis, Epicrius
Monachus
monachus, Monachus
monocarpa, Hydractinia
montanus, Peltops
montium, Paramythia
montium, Paramythia montium
montona, Campephaga .
montona, Edolisima montona .
morisii, Abraliopsis
moschata, Eledone
moschatum, Dicyemennea
multicornis, Perigonimus
multicristata, Vermiliopsis
multidentata, Goniada .
Muscipeta ....
muscoides, Bougainvillia
muscoides, Perigonimus .
muscus, Perigonimus
musseraensis, Diopatra .
Myiolestes ....
Myzomela ....
nanellus, Bougainvillia .
nanellus, Perigonimus
napolitanus, Perigonimus
261, 265,
356,
262,
351-359
261
337.
277, 279
67, 113
261, 270
• 3ii
• 301
44
. 422
• 425
• 399
68, 138
• 414
• 4*7
. 417
. 412
66, 97
432, 433
• 43i
• 433
• 317
185-188
39i, 423
• 423
424, 425
• 423
. 404
• 404
• 307
. 417
34L 347
. 341
. 188
358, 359
341, 347
277, 279
175-183
203-256
205-256
360-361
416
435
435
409
409
264, 279
283-291
291
341, 347
69, 145
67, 105
• 4i7
342, 346
342, 346
342, 346
67, 115
. 421
• 43i
342, 347
342, 347
342, 348
nasutus, Ophiopsiseps
367, 377
Neanthes ....
66
neapolitana, Diopatra
67, 116
Neopsittacus
402
Neositta ....
39i, 43o
Neoturris ....
• 338
Nephthys ....
67, 102
Nereis ....
66, 93
nigropectus, Eulacestoma
. 421
ninnetta, Lysidice
67, 114
Ninox .....
• 4°5
nordgaadi, Bougainvillia
• 342
niveifrons, Pachycephala rufinucha
. 420
norvegica, Hydroides
69, 143
Notostraca ....
• 3-57
nudus, Perigonimus
342, 347
nutans, Perigonimus
342, 347
obliqidens, Haplochromis
. 301
obscurior, Pachycephala schlegelii
. 420
occidentalis, Octopus
. 156
Oceania ....
• 344
ochrogaster, Alcyone azurea
39i, 408
octona, Leuckartiara
337-347
Octopoda ....
. 271
Octopodidae ....
. 272
Octopodoteuthidae
265
Octopus . . 260, 272, i
-79. 283-286
oculata, Phyllodoce
66, 85
Ocythoe ....
260, 274
Ocythoidae ....
. 274
officinalis, Sepia ... 2
60, 262, 276
Ommastrephes ... 2
61, 267, 268
Ommastrephidae .
267, 279
Onychia ....
261, 265
Onuphis ....
67, 118
Onychoteuthis
261, 266
Ophioglycera
67, 107
Ophioseps .... 3
65. 367. 368
Opisthosyllis ....
66, 90
Orbinia ....
67, 122
Oreopsittacus
402
Oreornis ....
• 434
Oreostruthus
. 436
ornata, Hydractinia
• 355
ornata, Psittacella brehmi
. 403
ovum, Buccinum .
• 358
Owenia ....
68, 131
oweniana, Sepietta . . 2
61, 263, 276
Pachycephala
420
packardi, Lepidurus apus
5i
palliata, Atractylis
• 342. 347
palliatus, Perigonimus
. 342
pallida, Psittacella brehmii
. 403
palustris, Porzana
• 399
palustris, Porzana pusilla
39i. 399
Pandeidae ....
• 347
Pandeinae ....
• 347
444
INDEX
Panthalis
65, 74
prampramensis, Trypanosyllis
66, 88
papanua, Microeca
. 417
prampramiana, Vermiliopsis .
69, 146
papuana, Microeca
. 417
princeps, Melidectes
. 433
papuensis, Podargus
. 406
Prionospio ....
67, 124
papuensis, Tyto longimembris
. 405
Proboscidactyla
• 339
Paradelma
. 369
Proboscidactylidae
• 347
Paradisaea
425
prodrumus, Haplochromis
305,
308, 311,
Paralabidochromis .
299-327-332
3i8, 331
Paramythia .
• 435
Pseudone 
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